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HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

BULLETIN

OF THE

MUSEUM OF COMPAEATIVE ZOOLOGY

HARVARD COLLEGE, IN CAMBRIDGE

VOL. 125

CAMBRIDGE, MASS., U.S.A.
1961

The Cosmos Press, Inc.
Cambridge, Mass., U.S.A.

CONTENTS

PAGE

No. 1. — Shell Morphology in the Larval and Post-
larval Stages of the Sea Scallop, Placopecten
magellanicus (Gmelin). By Arthur S. Merrill.
(3 plates.) March, 1961 ..... 1

No. 2. — On the Major Arterial Canals in the Bar-
Region of Testudinoid Turtles and the Classi-
fication of the Testudinoidea. By Samuel
Booker McDowell, Jr. June, 1961 .... 21

No. 3. — Two Jurassic Turtle Skulls : A Morphological
Study. By Thomas S. Parsons and Ernest E. Wil-
liams. (6 plates.) June, 1961 .... 41

No. 4. — On the Species of Pseudidae (Amphibia,
Anura). By J. M. Gallardo. (1 plate.) August,
1961 109

No. 5. — Correlations Between Ecology and Morphol-
ogy in Anoline Lizards from Havana, Cuba
and Southern Florida. By Bruce B. Collette.
August, 1961 135

No. 6. — Cranial Anatomy of the Cynodont Reptile
Thrinaxodon liorhinus. By Richard Estes. (2
plates.) August, 1961 163

No. 7. — Two Sympatric Cuban Anoles of the caro-
linensis Group. By Rodolfo Ruibal and Ernest
E. Williams. August, 1961 181

No. 8. — The Taxonomy of the Anolis iiomolechis
Complex of Cuba. By Rodolfo Ruibal and Ernest
E. Williams. August, 1961 209

PAGE

No. 9. — Taxonomy of the Deep Sea Fishes op the Genus
Chauliodus. By -James E. Morrow, Jr. October,
1961 . . ' 247

No. 10. — A Contribution to the Biology of the Gigan-

TURIDAE, WITH DESCRIPTION OF A NEW GENUS

and Species. By Vladimir Walters. October, 1961 295
No ii, — Variation in Paramyxine, with a Redescrip-

TION OF P. ATAMI DEAN AND P. SPRINGER! BlGELOW

and Schroeder. By Pi. Strahan and Y. Honma.
October, 1961 .' 321

No. 12. — Abyssal Mollusks from the South Atlantic
Ocean. By Arthur H. Clarke, Jr. (4 plates.)
October, 1*961 343

No. 13. — The Genus Micrathena (Araneae, Argiopidae)
in Central America. By Arthur M. Chiekering.
November, 1961 389

No. 14. — The Proscalopinae, a New Subfamily of Talpid
Insectivores. By Katheriue M. Reed. (2 plates.)
November, 1961* 471

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Volume 125, No. 1

SHELL MORPHOLOGY IN THE LARVAL

AND POSTLARVAL STAGES OF THE

SEA SCALLOP, Placopecten magellanicus (Gmelin)

By Arthur S. Merrill

WITH THREE PLATES

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

March, 1961

Publications Issued by or in Connection
with THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 - - The current volume is Vol. 125.

Breviora (octavo) 1952 — No. 134 is current.

Memoirs (quarto) 1864-1938 — Publication was terminated with
Vol. 55.

Johnsonia (quarto) 1941 — A publication of the Department of
Mollusks. Vol. 3, no. 39 is current.

Occasional Papers of the Department op Mollusks (octavo)
1945 — Vol. 2, no. 25 is current.

Proceedings op the New England Zoological Club (octavo)
1899-1948 — Published in connection with the Museum. Publication
terminated with Vol. 24.

The continuing publications are issued at irregular intervals in num-
bers which may be purchased separately. Prices and lists may be
obtained on application to the Director of the Museum of Comparative
Zoology, Cambridge 38. Massachusetts.

Of the Peters "Check List of Birds of the World," volumes 1-3 are
out of print ; volumes 4 and 6 may be obtained from the Harvard Uni-
versity Press; volumes 5, 7 and 9 are sold by the Museum, and future
volumes will be published under Museum auspices.

Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE

Volume 125, No. 1

SHELL MORPHOLOGY IN THE LARVAL

AND POSTLARVAL STAGES OF THE

SEA SCALLOP, Placopecten magellanicus (Gmelin)

By Arthur S. Merrill

WITH THREE PLATES

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

March, 1961

Xo i _ _ Shell Morphology in the Larval and Postlarval Stago
of the Sea Scallop, Placopecten magellanicus (Gmelin)

By Arthur S. Merrill1

CONTENTS

Page

Introduction and Acknowledgments 3

The Larval Shell 6

The Postlarval Shell . . 7

Shape and Color 7

Shell Structure 9

The External Sculpture 10

The Inner Shell Surface 1-

The Auricles or Wings 13

Hinge-Plate ... 1 !

The Byssal Notch 15

The Ligament 15

Discussion ... ' ( '

References 1 s

INTRODUCTION AND ACKNOWLEDGMENTS

The larval and postlarval stages of many species of pelecypods
have been studied and reported by various investigators over the
years. Stafford (1912) and Sullivan (1948), working with
species from the northeast coast of North America, have con-
tributed materially to our knowledge of several species. However,
the sea scallop, Placopecten magellanicus (Gmelin), has received
very little attention. Sullivan's study did not include the spe-
cies, and Stafford only devoted half a page to it. Consequently,
efforts to identify the early stages of the sea scallop by means of
the existing literature have met with little success.

Attempts at rearing larvae of the sea scallop have also been
unsuccessful (Drew, 1906; Posgay, 1953 ; Baird, 1953). There-
fore, it has been necessary to compare the changes and modifica-
tions through an extensive series of specimens in order to identify
the early stages of the species with certainty.

There is evidence that past investigators have experienced
difficulty in identifying the young of the sea scallop. Dall (1898,
p. 726) lists an excellent synonymy which includes several specific
names thought to be new species but which later were found to
be young sea scallops.

The primary purpose of this paper is to describe and figure

iU. S. Department of the Interior. Fish and Wildlife Service, Bureau of Com-
mercial Fisheries, Biological Laboratory, Woods Hole, Massachusetts.

4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

development of shell characters through the early stages of this
commercially important species with the hope that it will allow
future workers to quickly and accurately identify it at any stage
of growth. Descriptions of the adult stage are included when
it is necessary to describe later development of a particular
character.

Little has been written of the early life history of the sea
scallop. Jackson (1890) described and figured specimens he
thought to be young P. magellanicus but which proved to be
Cyclopecten nanus, a species subsequently described and figured
by Verrill in 1897 (see Merrill, 1959). Drew (1906) fully de-
scribed and figured both the shell and the animal of the adult,
but his attempts to rear the young were unsuccessful and he was
only able to discuss and figure the very early stages of em-
bryogeny covering the first 5 days after fertilization. Stafford
(1912) was able to identify Pecten larvae found in the plankton
collected in his area but I am not convinced he could identify
them with certainty below generic rank. His last paragraph
about scallops on page 239 is ambiguous. If I interpret cor-
rectly, he ends his discussion by concluding that the P. magel-
lanicus in his figure 35 is Pecten islandicus. In any case, he
based his identity of allied species of Pecten on the differences
in size of the settling larvae, but others (Jorgensen, 1946; Sul-
livan, 1948; Loosanoff and Davis, 1950) have found larval size
to be quite variable within a species. My own observations agree
with this. Morse (1919, p. 152, fig. C) does little more than figure,
without measurements, the general outlines of an early stage
of the sea scallop. The figure, as he says, resembles certain
avicular forms. Borden (1928) gives measurements of pecten
larvae, all under 0.2 mm., taken from plankton tows. However,
her description of the larval form at about 0.3 mm. is incorrect,
which suggests that the still younger larval forms under 0.2
mm. may not have been correctly identified. Baird (1953)
published an interesting paper on the finding of a large number
of tiny sea scallops on Bryozoa. However, his work is not de-
scriptive ; rather, he discusses the unsuccessful laboratory
attempts to rear sea scallops beyond the trochophore stage and
relates his findings to age and growth in small scallops. The
present author (Merrill, 1959) described and compared the
juvenile sea scallop with a somewhat similar appearing species.
Cyclopecten nanus Verrill, but found it beyond the scope of that
paper to describe and figure the various early stages of shell
morphology in the sea scallop.

MERRILL : SHELL MORPHOLOGY OF PLACOPECTEN 5

The material used in this study came from several sources.
Sea scallops were reared in this laboratory at Woods Hole,
Massachusetts, through the straight-hinge stage. A large sample
of minute sea scallops was collected from a navigation buoy
brought in to the Coast Guard Station at Woods Hole, Massa-
chusetts, in June 1958. Nearly 10,000 of these tiny shells were
measured, of which 80 per cent were under 4 mm., the smallest
0.5 mm., the largest 13.2 mm. Several samples were taken from
other buoys as well as from the ocean bottom during the year
1958. In these samples, sea scallops as small as 0.25 mm. were
collected. This is about the size at spatfall when the scallops are
undergoing metamorphosis. Samples of large sea scallops taken
from various stations along the coast were also available for
study. The author also had ample opportunity to observe and
compare the structures and variations not only in these but in
other small scallops accumulated over the years from several
sources by personnel of the Bureau of Commercial Fisheries.
Biological Laboratory, at Woods Hole, Massachusetts.

The methods used in cleaning the minute completed larval
shells, and in separating and mounting the valves are those de-
scribed by Rees (1950, p. 75).

The term height refers to the distance from the umbo to the
ventral margin ; length to the distance from the anterior to the
posterior edge (see Fig 1) .

Acknowledgments are due the following individuals without
whose help and cooperation the value of this work would have
been considerably lessened. To Mr. Julius A. Posgay and Doctors
Robert L. Edwards and Roland L. Wigley of the Bureau of Com-
mercial Fisheries, Biological Laboratory, at Woods Hole, Massa-
chusetts, and to Doctors William J. Clench and Ruth D. Turner
and Mr. Robert Robertson of the Mollusk Department in the
Museum of Comparative Zoology at Harvard University, I am
greatly indebted for advice and comments after reading the
manuscript. To Mr. Robert Brigham, staff photographer of the
Bureau of Commercial Fisheries, Biological Laboratory, Woods
Hole, and Mr. Frank AVhite, staff photographer in the Biological
Laboratories at Harvard University, I owe much for the quality
of the photographs reproduced herein. I am also grateful
to Mr. Frank Bailey of the Bureau of Commercial Fisheries.
Biological Laboratory, at Woods Hole for the excellent line draw-
ings.

b BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

THE LARVAL SHELL

A prodissoconch or first-formed larval shell, strikingly dif-
ferent from the succeeding dissoconch, is easily seen in young post-
larval specimens of P. magellanicus (PI. 2, fig. 1). The pro-
dissoconch is distinctly separated from the dissoconch by a sharp
line. This heavy concentric line denotes the end of the free-
swimming pelagic period of the tiny scallop.

The larval development is usually divided into two stages.
The first is the "straight-hinge stage" in which the veliger pro-
duces the embryonic "D-shaped" shell. This is followed by the
"umbo stage" in which the shape, texture, and hinge structure
are sufficiently differentiated to allow identification.

The straight-hinge larva of the sea scallop is similar to that
of other lamellibranchs. It has a wide hinge-line, is relatively
long in proportion to its height, and is almost equally rounded
at both ends (PL 1 ) . The two valves are hinged dorsally, the mar-
gins meeting ventrally and laterally. The minuteness of the shell,
its transparency, and lack of structure leave few characters
of importance to aid in identification at this early stage. Both
valves are similar in size, shape, and curvature. They are fairly
flat, and lack color. The hinge-line is usually straight, but in
occasional specimens, it curves inward slightly. The size ranges
from 60 to 80 microns in height and 80 to 115 microns in length
in 7 -day-old laboratory -reared specimens.

At the umbo stage, as seen in the completed prodissoconch,
the valves are discoidal, inflated, and nearly equal in size, shape
and curvature, the upper valve being slightly more convex. The
structure of both valves is homogeneous, opaque, non-prismatic,
and with exceedingly fine lines of growth etching the surface. The
anterior end slopes somewhat more steeply than the posterior end
causing the curvature to appear broadly rounded in the ventro-
posterior region. The hinge-line is rather wide and straight. The
provinculum is thin, raised, and lined with two rows of indistinct,
minute, transverse tooth-like projections which fade and disap-
pear on each side at some distance from the resilium (Fig. 1).
The umbos are elevated, subcorneal, and while occupying the
mid-dorsal portion of the valves, slant rearward to overlie the
hinge-line. The left umbo is the more prominent. The size

MERRILL : SHELL MORPHOLOGY OF PLACOPECTEN (

(height) ranges from 0.22 to 0.32 mm. at the completion of tlie
prodissoconch. The color is pale yellow to flaxen.

LENGTH

i
o

Figure 1. Line drawing of right valve showing dissoconch structure. In
ternal view. Height 0.5 mm.

THE POSTLARVAL SHELL

The changeover from a planktotrophic larval existence to a
more or less stationary juvenile and adult life necessitates marked
changes in the anatomy and shell formation of the sea scallop.
These changes, modifications, and additions in the form and
structure as seen at various stages in the progressive development
of the shell can be brought out more clearly by a systematic
description of each major character.

Shape and Color. The valves soon lose the equal convexity of
the larval form, the top valve retaining a similar convexity while
the bottom valve flattens considerably. The margins meet evenly
until the young postlarval stages reach about 5 mm. in height
when the valves begin to gape slightly but noticeably at each end
in the dorso-lateral region. The gaping increases with size and
is quite pronounced in adult specimens. The ventral margin is
thin and simple with the upppr valve slightly overlapping in

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

young specimens (Fig. 2). This feature remains constant during
development except in some old specimens in which the edge
thickens.

Figure 2. Line drawing of n specimen 17.2 mm. to show various structures
of the shell in cross-section.

The form of the shell changes with growth. The prodissoconch
is broadly rounded, slightly longer than high. The scallop con-
tinues to be slightly compressed dorso-ventrally until it reaches
a size of 1 to 1.5 mm. at which time the height and length are
about equal. The shell now gradually becomes more and more
compressed laterally so that it is at its greatest extent in specimens
between 70 and 90 mm. in height. Then the shell slowly begins to
broaden again, and specimens 110 to 120 mm. assume a broadly
rounded shape of similar height and length. The broadening of
the shell continues and, in an average series, specimens larger
than 120 mm. become increasingly longer than high. Individual
specimens sometimes vary considerably but the measurements in
Table 1 typify the general average of the sea scallop.

Table 1 - - Ratios of height to length illustrating the changes
in shell symmetry during growth

Height
(mm.)

Length
(mm.)

Ratio

Height
(mm.)

Length
(mm.)

Ratio

*0.072

0.088

1.22

78.3

73.1

0.93

**0.30

0.32

1.07

96.5

92.5

0.96

0.55

0.57

1.04

112.2

1.00

1.32

1.00

122.0

124.2

1.02

2.30

2.16

0.94

140.8

146.4

1.04

10.0

9.2

0.92

178.0

188.0

1.06

58.1

52.8

0.91

***208.0

230.0

1.11

*Str;tight-hinge veliger
* "Completed dissoconch
•••Largest scallop recorded in the literature (See Norton 1931)

MERRILL : SHELL MORPHOLOGY OP PLACOPECTEN 9

Color pigment is lacking in the early dissoconch but when it
reaches about 1.5 mm. a light cream tint develops. From this
size on, color of various intensities ranging from light brown to
dark reddish brown becomes increasingly prominent. Occa-
sionally, albino, yellow, lavender, or radially dark- and light-
rayed specimens are produced. Sometimes the color changes as
the shell increases in size. Brown or reddish brown is the usual
adult color. The under valve usually remains white or cream-
colored throughout growth.

Shell Structure. The prodissoconch is thin and hyaline and con-
tinues translucent through the first few millimeters of growth of
the postlarval shell. There is a complete change in the structure
of the shell of the dissoconch. The larval shell with its high
cuticular content is succeeded by the postlarval shell containing
a high percentage of calcium salts. The shell is no longer simple;
rather it becomes layered and more complex in structure.

As is well known, the periostracum, shell and ligament are
products of secretion from the mantle. Outer and inner shell
layers are formed in the dissoconch of Placopecten with color
pigment being deposited in the outer layer. The periostracum
and the outer shell layer are secreted by the lobes of the mantle
edge with a resulting increment at the periphery. The general
surface of the mantle is responsible for the secretion of the inner
layer, the amount an I regularity of which subsequently deter-
mines the shell tbickness. The layers begin to form from the
moment dissoconch growth commences. An additional layer, dif-
ferent in structure and composition, is formed by the modified
mantle epithelium of the adductor muscle. This layer is seen to
fracture on a vertical plane with respect to the shell surface.
Tt is difficult to make out any microscopic features in this layer
without using special techniques because of the homogeneity and
transparency of the substance. In Pec+en, according to Boggild
(1930), this layer is prismatic in structure and the prisms are
very thin, straight, regular, and oriented to a vertical axis.
Boggild found (using certain techniques of refraction) that this
layer is aragonitic as opposed to the calcific outer and inner layers.
The layer is first apparent in the sea scallop at about 2 mm. The
outer layer is thin and only clearly visible in sectioned shells 15
mm. or less in size while the inner layer is much thicker and can
be seen in all sbes. The inner layer is built up of irregular
parallel leaves to appear as a foliated structure.

Figure 2 shows a transverse section of a juvenile sea scallop

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

of 17.2 mm., drawn with the aid of a camera lucida. Of interest
is the relative thickness of the valves, the amount of overlap of
the upper valve at the ventral margin, the position of the upper
and lower muscle scars, and the arrangement of the hinge and
ligament. Of particular interest is the method of development
of the layer produced by the muscle epithelium. The adductor
muscle must necessarily change position as the shell enlarges in
order to maintain a somewhat central location. This is accom-
plished by a ventral migration of the muscle. At the same time
deposition of shell material continues in the area contiguent to
the muscle so that in time a distinct middle layer dorsal to the
muscle is produced.

A well-defined prismatic layer is easily seen in the lower valve
of the young dissoconch when viewed from above (Figs. 1 and 3;
PI. 2, fig. 2). The prisms are irregularly shaped and are secreted

0. I mm

-H

Figure 3. Line drawing to show prismatic structure in the right valve.

in an erratic, though concentric pattern. On the average, when
measured near and parallel to the ventral margin, there are
about 5 prisms per 0.1 mm. The prismatic pattern persists until
the shell reaches about 5 to 7 mm. in height at which size the
shell becomes too thick for this structure to be seen clearly.
The External Sculpture. The surface sculpture differs in the

MERRILL: SHELL MORPHOLOGY OF PLACOPECTEN 11

two valves. In the upper valve, commencing with the onset of
dissoconch growth, microscopic radial ridges develop which are
crossed at intervals by similarly delicate concentric ridges which
result in a sharply defined cancellated or nodulated sculpture.
At about 2 or 3 mm. the concentric ridges weaken and gradually
disappear whereas the fine radial ridges are retained. Shortly
thereafter heavier raised primary ribs begin to radiate regularly
among the intervening microscopic ridges which in turn fade
and cease to be produced in juvenile specimens hetween 10 and
20 mm. The fine ridges may be straight and regular or they may
be vermicular or oblique in which case they usually cross the
ribs. This is the "camptonectes" sculpture of Verrill (1897).
Commencing with the juvenile, the regularly spaced primary ribs
are a prominent part of the upper valve, new ribs being added
between them as the shell enlarges (PI. 3). When they first
appear, the ribs are spaced about 0.3 mm. apart. When the spac-
ing increases to 0.8 to 1.2 mm., a new rib is added. The distance
between the ribs increases as the shell size increases until in the
adult stage the ribs are spaced about 1 mm. apart. The ribs of
the adult may be strongly raised and serrated causing the shell
to look and feel rough, or, in the case of a smooth-shelled adult,
the ribs may be but slightly raised.

Any concentric markings beyond that mentioned for the early
dissoconch are in the form of numerous fine laminal lines (striae)
as the result of frequent additions during growth. At intervals
these lines of growth become more closely spaced appearing as
thickened bands of shell material. Stevenson and Dickie (1954)
and most present-day investigators working with the sea scallop
believe that the more prominent bands are caused by the seasonal
changes of growth pattern, as has been demonstrated with the
annual rings of trees or fish scales.

Radial sculpture is formed somewhat similarly in the bottom
valve. However, due to the formation of prismatic structure in
the early dissoconch the fine ridges do not start forming until
the shell is about 5 to 7 mm. in height. Primary ribs soon develop
as in the upper valve but not as strongly. Although both the
ribs and ridges usually persist through to the adult, either or
both may disappear during the course of development. If the
sculpture persists it is weaker than in the upper valve.

The radial sculpture of the wings is similar to that on the sur-
face of the shell, but it is usually more strongly developed, espe-
cially so in the anterior wings (PI. 3). Concentric sculpture

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ending at the wing margins forms crenulations at the hinge line.
The Inner Shell Surface. The mantle is attached to the shell
by a series of muscles alonir the pallial line. The mantle is ahle to
expand and retract from this line to the outer edge of the shell. In
the adult the surface area outside the pallial line appeal's smooth
and lustrous to the eye but upon closer examination flakey laminal

Figure 4. A series of line drawings showing the development of the wings
of the sea scallop.

layering can be observed, particularly near the margin. Also,
very tine lines radiate to the margin. Within the pallial line, the
calcium deposits appear nacreous with coarse and irregular lay-
ering. The adductor muscle scar is large, distinct, and, partic-
ularly on the right valve, placed off center toward the posterior
border. The scar on the left (upper) valve is the larger and
there is a line showing the separation of the muscle into a large
anterior and a small posterior part. Within the scar, lines de-
noting the attachment of individual bundles are evident. The
adductor muscle is attached to the valves somewhat more dorsallv

MERRILL: SHELL MORPHOLOGY OF PLACOPECTEN 13

and posteriorly in the young especially in the right valve, but it
gradually moves to a more central location with increase in shell
size, and after about 50 mm. advances proportionally to the ven-
tral margin of the shell. The muscle scar forms a slight depres-
sion and is surrounded by the crystalline substance. The peculiar
crystalline structure within the pallial line is first evident in the
upper valve at about 2 mm. but soon can also be seen in the
thinner bottom valve.

The Auricles or Wings. Figure 4 compares the outline of speci-
mens in a series ranging in size from 0.3 to 16.5 mm. in height.
These were specifically drawn to show the development of the
wings. As can be seen, the settled prodissoconch lacks them. The
earliest indication of their development appears in specimens
which have put on just a fraction of a millimeter of dissoconch
growth. A notch develops in the lower valve at the anterior
margin, the portion of the newly-formed shell dorsal to the notch
consequently appearing ear-like. The wing becomes broadly
rounded as the height increases to about 2 mm. It now begins
to angulate (Fig. 4b) and continues until the shell is about 10
mm. in height when the angle again broadens (Fig. 4c). The
angle becomes less acute as the shell size increases so that in the
adult this wing is more nearly symmetrical with the posterior one.

The development of the anterior wing in the upper valve pro-
ceeds in a peculiar fashion. A rounded flap begins to develop
almost immediately, projecting over and overlying the byssal
notch. Although fully developed at 1 mm., it persists through
growth to nearly 5 mm.

At the same time as these wing-like processes are developing,
slight indentations or depr ssions begin to form between them
and the body wall. These are the slopes or submargins, the points
where the wings can be distinguished from the body proper.

The posterior wings begin to develop at about 2 mm. and, with
increase in size, slowly become more strongly outlined as the
posterior submargins strengthen. The wings are easily distinguish-
able before 5 mm., becoming even better developed as the shell
size increases. However, it is interesting to note that the ears
are proportionally smaller in larger specimens. At 2 mm. the
width of the hinge-line is about four-fifths the length of the shell,
at about 35 mm. only about one half, and thereafter very gradu-
ally decreasing. The shell measurements in Table 2 clearly show
this characteristic.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Table 2 - - Width of hinge-line compared to length showing the
decrease in the wing width in relation to length with growth

(in millimeters)

Width of Hinge-line „„

Height
1.9

Length

1.8

Width of hinge-line

1.5

Length

83.3

5.8

5.2

3.8

73.1

10.5

9.1

6.3

69.2

21.3

18.9

11.6

61.4

34.6

31.6

16.2

51.3

52.3

48.0

20.8

43.3

106.8

104.2

44.3

42.5

142.6

147.7

62.0

42.0

Ilintj) -Plate. In young specimens the hinge-plate is thin, raised,
and with a simple ligamental groove below the margin which is
separated by an internal resilium (Fig. 5). The indistinct trans-
verse teeth present in the prodissoconch cease to develop in the

Figure 5. Line drawing of right valve of a specimen 5.7 mm. showing
hinge-line and other internal characters.

early dissoconch. As the hinge-line develops, deposition of new
material obliterates the larval teeth at about 0.5 mm. The hinge-
plate, only slightly differentiated in the very young, becomes
increasingly raised along the exterior margin as the shell develops

MERRILL : SHELL MORPHOLOGY OF PLACOPECTEN 15

and this is directly responsible for the greater gaping of the
valves in the dorsolateral region. At the hinge-line, the exterior
margin is creimlated and recurves inward to impinge on the
hinge-plate and partly overlap the ligamental groove. This be-
comes more apparent with increased shell size.

The Byssal Notch. Formation of the byssal notch commences
at the moment of dissoconch growth. In the early dissoconch,
little shell material is deposited in the lateral region of the under
valve where the notch is located ; instead, a rounded indentation
is produced. With increase in size the notch advances accordingly
as shell material is deposited behind it (Fig. 1). This material
is not prismatic as is the rest of the valve in the early dissoconch ;
rather it appears homogeneous and hyaline, with laminated, con-
centric growth lines running parallel to the notch. As the shell
enlarges and the notch continues to advance, a ridge is produced
along the anterior submargin above which a groove has been
excavated. A number of pectinidial teeth develop along the ridge.
Teeth begin to form at a size of about 1.5 mm. (PI. 2, fig. 2) and
cease after about 25 mm. Occasionally, the teeth are feeble or
may not develop at all. In one specimen over 30 teeth were
counted. Plate 3, figure 1 illustrates a 5.7 mm. specimen in
which 15 teeth are present. The early-formed teeth can usually
he observed in the adult well back on the raised ridge in the
vicinity of the umbonal region unless erosion or calcification has
destroyed or covered them. The notch is broad and rounded inside
for the first millimeter or two (Fig. 1) but soon begins to angu-
late as the wing develops. After a size of 10-15 mm. the angle
becomes gradually less acute (PI. 3, fig. 3) as the notch fills in so
that in the adult the notch is almost entirely obliterated.

The L'gami nt. The structure, function and mechanical prop-
erties of the ligament in reef en have been fully described and
discussed by Trueman (1953a, b). Briefly, the ligament joins
the two valves of the shell together dorsally along a hinge-line
and functions by working in opposition to the adductor muscle,
thus forcing the valves to gape apart when the muscle is relaxed.
There are two layers, the outer which extends thinly along the
dorsal margin of the valve hinges, and the inner which is centered
immediately below the hinge at the umbo. A thin layer of perio-
stracum is produced over the outer layer. The layers are lamin-
ated and, according to Trueman (1953a), composed largely of
tanned proteins, an organic matrix commonly referred to as
"eonchiolin."

16 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

The inner layer, or resilium, is triangulate in shape, its apex
coincident with the umbonal point of the valves. It is housed in
a depression, the resilifer, the base of which projects shelf-like
into the cavity of the shell. It is composed of three parts : a large
central region consisting of amorphous, non-calcareous tanned
proteins, brown in color and of a rubbery consistency, and two
small lateral calcified regions in the area of attachment to the
resilifer.

In the early postlarval shell of the sea scallop the outer liga-
ment lines the hinge as a very thin, transparent substance. As
the shell enlarges this layer becomes increasingly conspicuous
principally because the brown color in it is darker.

In the larval shell of Pecten the ligament is situated internally
and off center on the straight hinge toward the posterior broad
end of the larva (Rees, 1950). In the recently settled larva of
the sea scallop the resilium is about centered between the umbos
(Pig. 1). It is wide and rounded, and seated within a weakly
developed resilifer. After the first millimeter of growth the re-
silium begins to lengthen and soon takes an almost equal-sided
pyramidal shape as new ligamental material is added ventrally
and laterally. After 5 mm. (Fig. 5) to 10 mm. the resilium grad-
ually begins to elongate in a dorso-ventral direction until in the
adult it is long and slender. The resilifer strengthens with
growth, the depression gradually deepening so that in the adult
the resilium is deeply rooted.

The ligament, formed by secretions from the mantle, is subject
to the same general growth and shock lines as is the shell. Less
prominent impressions are found in the tanned protein portion
than in the calcified outer portion of the inner ligament.

DISCUSSION

The major changes during the development of the shell of the
sea scallop have been described in the preceding pages, thus serv-
ing the primary purpose of this paper. However, the subject
would not be complete without a brief discussion of certain points
described elsewhere.

The D-shaped veliger shell is usually distinctly marked off
from the umbo veliger stage in most species (Rees 1950). This
is not the case with the sea scallop. Apparently there is little or
no cessation of growth during larval shell development to cause
this distinct line to be formed. Thus, the early veliger is almost
impossible to identify in the plankton. Also the size at this stage

MERRILL : SHELL MORPHOLOGY OF PLACOPECTEN 17

varies considerably and is of little use as a character. This agrees
with Jorgensen's (1946) findings regarding the reliability of the
length of Prodissoconch I ["straight-hinge" stage] as a specific
criterion. Of course, by the time of development to the umbo
stage this species should be identifiable through hinge characters.
Stafford (1912) states that larval sea scallops are easily identi-
fied because of their equal measurements in length and depth.
Of the scallops examined in this study, the length commonly
exceeded the height in the prodissoconch. Stafford also speaks of
a notch that may appear in front of the median point of the
margin, which soon develops into two little beaks on the left
valve, and a larger interlocking tooth on the right valve. Borden
(1928) also remarks on this occurrence but it is quite possible
she simply followed Stafford's earlier work. No such anomiaiid-
like character was observed in any of my specimens. In fact, I
should be very much surprised if snch a character existed in this
species, principally because it is not a known phylogenetic char-
acter in the Pectin idae.

The prismatic structure in the under valve of the early post-
larval sea scallop has been described elsewhere in this paper. In
very young specimens the margin of this valve is very thin and
flexible due to the organic matrix bordering each prism. Similar
prismatic structure can be seen in other species of the Pectinidae,
and is especially prominent in thin-shelled deep-water forms.
Carpenter (1847, p. 95) first observed this character in Pccten
nobilis. The flexibility of the under valve allows the shells to com-
press snugly together when necessary and seems to serve essen-
tially to give extra protection.

The very young sea scallop puts out a short branching byssus
which allows it to adhere closely to the surface to which it
attaches. The rounded flap that precedes the deve'opment of the
anterior wing overlies the byssal notch and appears to function
as a protective lid in the young dissoconch. Thus when a young
scallop is disturbed it squeezes its valves tightly together and, be-
ing attached close to the surface with its attachment protected, is
better able to withstand and survive the early vicissitudes of life.

Before leaving the subject of byssal formation, it is of interest
to note that the sea scallop retains the habit of byssal fixation for
some time after reaching adult size. T have measurements of
scallops as large as 112 mm. in height which were found still
attached to rocks. The scallop does not remain attached all of

18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

this time. It frequently breaks away and swims about for a
time before attaching again.

It should be noted that Dall (1886, pp. 207, 222) and Verrill
(1897, p. 79) both described well-developed transverse incisions
on the hinge-plate in young sea scallops as did Jackson (1890).
I have not been able to detect any signs of transverse grooves or
ridges in this species other than the very indistinct transverse
tooth-like projections in the larval shell which fade and disap-
pear very shortly in the dissoconch and which are invisible unless
the specimen is properly prepared and mounted. Apparently
the above authors, like Jackson, had the young of other species
before them when they described this feature.

Dall (1889) discusses the radiating ridges or ribs found in
certain groups of pelecypods and concludes that they serve to
add strength to the shell while not increasing the weight as would
a corresponding increase in thickness of the shell. As it happens,
the adult sea scallop increases the thickness of the shell rather
than develop strong radial sculpture. However, the young scallop
of 5 to 30 mm. (PI. 3) has rather strong ribs which would tend to
strengthen the shell before it has had a chance to thicken
markedly. The radiating ribbed sculpture of the wings is par-
ticularly strong. The wings at these sizes are proportionally
wider at this early age but thin, and the extra strength is desir-
able in this area of the pivotal axis of the valves where there are
mechanical stresses and reactions of one valve upon the other for
the scallop to contend with.

REFERENCES

Baird, Frederick T., Jr.

1953. Observations on the early life history of the giant scallop
(Pecten magellanicus). Bull. Dept. Sea and Shore Fish., Res.
Bull. no. 14, pp. 2-7.

BOGGILD, O. B.

1930. The shell structure of the niollusks. D. Kgl. Danske Vidensk.
Selsk. Skr., Naturvidensk. og Mathem. Afd., Kobenhaven, ser.
9, vol. 11, no. 2, pp. 233-325.
Borden, Mabel A.

1928. A contribution to the study of the giant scallop, Placopecten
grandis (S.). Fish. Res. Bd. Canada, MS. Rep. Biol. Sta., no.
350, pp. 1-35.
Carpenter, W. B.

1847. On the microscopical structure of shells. Rep. Brit. Assoc. Adv.
Sei., Seventeenth Meeting, Oxford, 1847, pp. 93-13«.

MERRILL : SHELL MORPHOLOGY OF PLACOPECTEN 19

Dall, William H.

1886. Report on the Mollusca. Part 1 — Branchiopoda and Pelecypoda.
Bull. Mus. Comp. Zool., vol. 12, no. 6, pp. 171-318.

1889. On the hinge of pelecypods and its development, with an attempt
toward a better subdivision of the group. Ainer. Journ. Sci.,
ser. 3, vol. 38, pp. 445-462.

1898. Contributions to the Tertiary fauna of Florida. Trans. Wagner
Free Inst. Sci. Philadelphia, vol. 3, pt. 4, pp. 571-947.
Drew, Oilman A.

1906. The habits, anatomy, and embryology of the giant scallop (Pec-
ten tenuicostatus, Mighels). Univ. Maine Studies, no. 6, pp. 3-71.
Jackson, Robert T.

1890. Phylogeny of the Pelecypoda, the Aviculidae and their allies.
Mem. Boston Soc. Nat. Hist., vol. 4, no. 8, pp. 277-400.

Jorgenson, C. Barker

1946. 9. Lamellibranchia. Reproduction and larval development of
Danish marine bottom invertebrates. Medd. Komm. Havunder-
sog., Kobenhaven, Series: Plankton, vol. 4, no. 1, pp. 277-311.
Loosanoff, Victor L. and Harry C. Davis

1950. Conditioning V. mercenaria for spawning in winter and breeding
its larvae in the laboratory. Biol. Bull., vol. 98, no. 1, pp. 60-65.
Merrill, Arthur S.

1959. A comparison of Cyclopecten nanus Verrill and Bush and Placo-
pecten magellanicus (Gmelin). Occ. Papers on Mollusks, Har-
vard Univ., vol. 2, no. 25, pp. 209-228.
Morse, Edward S.

1919. No. 5 — Observations on living lamellibranchs of New England.
Proe. Boston Soc. Nat. Hist., vol. 35, no. 5, pp. 139-196.
Norton, A. H.

1931. Size of the giant scallop (Pecten grandis Sol., P. magellanicus
GmeU. The Nautilus, vol. 44, no. 3, pp. 99-100.

POSGAY, J. A.

1953. Sea scallop investigations. Sixth report on the investigations of
the shellfisheries of Massachusetts. Div. Mar. Fish., Dept. Nat.
Res., Commonwealth of Massachusetts, Boston, pp. 9-24.

Rees, C. B.

1950. The identification and classification of lamellibranch larvae.
Hull Bull. Mar. Ecol., vol. 3, no. 19, pp. 73 104.
Stafford, Joseph

1912. On the recognition of bivalve larvae in plankton collections.
Cont. Canadian Biol. 1906-1910, no. 14, pp. 221-442.
Stevenson, J. A. and L. M. Dickie

1954. Annual growth rings and rate of growth of the giant scallop,
Placopecten magellanicus (Gmelin) in the Digby area of the
Bay of Fundy. Journ. Fish. Res. Bd. Canada, vol. 11, no. 5,
pp. 660-671.

20 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Sullivan, Charlotte M.

1948. Bivalve larvae of Malpeque Bay, P.E.I. Fish. Bes. Bd. Canada,
Bull. no. 77, pp. 1-36.
Trueman, E. R.

1953a. The ligament of Pecten. Quart. .Tourn. Micr. Sci., vol. 94, no. 2,

pp. 193-202.
1953b. Observations on certain mechanical properties of the ligament
of Pecten. Journ. Exp. Biol., vol. 30, no. 4, pp. 453-467.
Verrill, A. E.

1897. A study of the family Pectinidae, with a revision of the genera
and subgenera. Trans. Connecticut Acad., vol. 10, pp. 41-95.

PLATES

PLATE I
L'igurc 1. Seven-day-old laboratory-reared sea scallop in the straight
hinge stage. Animal within shell (about 300X).

Figure 2. Same as Figure 1. Shell valves only (about 500X).

PLATE 2
Figure 1. Right (bottom) valve of early postlarval sea scallop. Heighl
0.5 mm.

Figure 2. Right valve showing first pectinidial tooth. Height 1.5 mm.

PLATE 3

Figure 1. Left and right valve of a specimen 5.7 mm. in height (holli
valves doubled).

Figure 2. Left and right valve of a specimen 11.7 mm. in height.
Figure 3. Left and right valve of a specimen 20.5 mm. in height.

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Bulletin of the Museum of Comparative Zoology

AT HAEVAKD COLLEGE

Vol. 125, No. 2

ON THE MAJOR ARTERIAL CANALS

IN THE EAR-REGION OF TESTUDINOID TURTLES

AND THE CLASSIFICATION OF THE

TESTUDINOIDEA

By Samuel Booker McDowell, Jr.

Newark College, Rutgers the State University, Newark, New Jersey.

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

June, 1901

No. 2 — On the major arterial canals in the ear-region of
Testudinoid Turtles and the classification of the Testudino'dea

By Samuel Booker McDowell, Jr.

That there is an osteological difference in cranial foramina
between typical emydine and testudinine turtles, on the one hand,
and kinosternines (mud-turtles), on the other, has been known
for some time. Siebenrock illustrated (1897, pi. 2, fig. 8) the inner
view of the cranial chamber of "Cinosternum odoratnm"
(=Sternotherus odoratus) and showed a foramen labeled f.c.i/
=" Foramen caroticum internum, Zweig zur Augenhohle hinzie-
hend" that does not appear in his figures of other Testudinoidea
(Chelydra serpentina, Testuno (sic, =Psammobatcs) tentoria,
Cyclemys dhor (=C. dentata), and Geoemyda spinosa). I can find
no discussion in Siebenrock 's text on this foramen in "Cinoster-
num."

A more striking difference, because it is more easily observed,
was still earlier noted by Baur (1888b), who found that an
arterial foramen between the pro-otic and quadrate seen in nearly
all turtles is greatly reduced or absent in Dermatemys, Stauroty-
pus, and the kinosternines. This feature was confirmed by Bienz
(1895) and is cited, as one of many osteological characters, in
the classification of turtles by Romer (1956) .

But while this characteristic has long been known in terms of
osteology, there is no work, to my knowledge, describing the dif-
ference between mud-turtles and pond-turtles in the arterial
supply of the head. Since closure of a large arterial foramen
implies considerable rearrangement of the head arteries, a
knowledge of the arteries would seem essential to interpretation of
the osteological features.

Dissection of Kinosternon subrubrum and Sternotherus odo-
ratus (kinosternid foramen-pattern) and of Malaclemys (Grap-
temys) geographica, Chrysemys (Trachemys) scripta, Chrysemys
picta, Gopherus berlandieri, Testudo (Testudo) graeca, Ocadia
sinensis, Chinemys reevesi, Clemmys inscidpta, Emys orbicularis
(Kunkel's series F slides, Kunkel, 1912), and Chelydra serpenti-
na (juvenile) (testudinid foramen pattern) convinces me that
the two foramen patterns are indicative of two quite different
adult arterial patterns, and further, that neither arterial pattern

24 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

can be derived directly from the other one, although both may
be derived from a more generalized pattern, such as that seen
in living Cheloniidae.

I. THE PATTERN OF THE TESTUDINIDAE

Bony canals and foramina: — There is an opening on the rear
surface of the skull, called the posterior aperture by some authors
= the foramen jugulare internum of Siebenrock (1897), the
fenestra postotica of Nick (1913) . This opening is bounded later-
ally by the quadrate, ventrally by the pterygoid, and medio-dor-
sally by a vertical lamina of the opisthotic or the exoccipital or
the two bones together. (Since this lamina represents a posterior
protection for the peculiar pericapsular extension of the perilym-
phatic sinus characteristic of turtles, I propose to call it the crista
postperilymphatica, new term.) Nick's name for the opening,
fenestra postotica, seems more fortunate than Siebenrock 's name,
foramen jugulare internum, since the opening rarely contains
the strict homologue of the mammalian vena jugularis interna.
Rather, it contains the main reptilian head-vein, the vena capitis
lateralis, sometimes called, by analogy, the "internal jugular,"
but more nearly homologous to the mammalian sinus cavernosus.
The turtle homologue of the mammalian vena jugularis interna is
a rather small vein closely bound to the vagus nerve, and with
the nerve piercing the crista postperilymphatica by an opening
I here call the apertura vagi, new term.1 The vein and vagus may
then be followed deeper within the skull and be seen to enter the
true foramen vagi or foramen jugulare between opisthotic and
exoccipital. The vena jugularis interna then extends mediad in
the dura mater above the brain (cf. mammalian sinus transver-
sa) to join a longitudinal vein in the dura mater on the midline
above the brain (cf. mammalian sinus sagittalis superior). In

^Throughout this paper in dealing with openings in the dried skull for cranial
nerves, I have used the word "foramen" for holes in the primary braincase, and
the word "apertura" for holes in the secondary investments of the primary
braincase. This convention is particularly necessary in dealing with holes for
transit of the glossopharyngeal nerve ; in such turtles as Orlitia this nerve
pierces hone three times: (1) an internal glossopharyngeal foramen leading from
the cranial cavity into the inner ear cavity of the dried skull, (2) an external
glossopharyngeal foramen from the inner ear capsule to the pericapsular space,
and (") a glossopharyngeal aperture in the crista postperilymphatica (a secondary
investment of the primary braincase). The second hole is invariably present in
turtles ; the first is often undefined by bone in young turtles, but is probably
constant for all fully mature skulls. The third hole is absent in adults of many
turtles. Without a convention the name "external glossopharyngeal foramen"
would be ambiguous and might refer either to the constant foramen in the
posterior wall of the ear capsule or to the variable "aperture" in the crista
postperilymphatica.

MCDOWELL: ARTERIES OF TESTUDINOID SKULLS 25

Platystcmon the apertura vagi is not defined laterally by bone
from the fenestra postotica, but is merely a deep notch, while in
Hardella it is a rather shallow notch. In other testudinoids,
however, the opening for the true internal jugular is set off by
bone from the fenestra postotica.

The fenestra postotica is usually triangular in form, and we
may speak of a "dorsal corner," a ventro-lateral corner," and
a "ventro-medial corner." In addition, the columella auris may
be seen passing across the fenestra as it extends from the fenestra
ovalis to the ear-chamber in the quadrate. Thus, it is sometimes
convenient to refer to a "supraeolumellar portion of the fenestra
postotica" and an "infracolumellar portion of the fenestra pos-
totica." The infracolumellar portion is a rather narrow horizon-
tal slit (for the only structures to pass here are the internal
carotid artery and the chorda tympani nerve and Jacobson 's and
the auricular anastomotic nerves) .

Deep within the skull, the supraeolumellar portion of the
fenestra postotica appears divided into two passageways. The
smaller passageway is at the dorsal corner and leads upward and
forward between the pro-otic and quadrate bones to open by a
foramen into the temporal fossa. This passageway and the fora-
men forming its opening to the temporal fossa are, respectively,
the canalis caroticus externus and foramen carotico-temporale of
Siebenrock (1897), while Nick terms the opening "Foramen
arteriae facialis, ' ' in accordance with a different terminology for
the vessel. Actually, the foramen transmits both the temporal
stem of the stapedial artery and a branch of the vena capitis
lateralis. Since the homology between the stapedial artery and
adult human external carotid is far from exact, the differences
being in precisely this region, I modify Siebenrock 's names to
canalis stapediarterialis and foramen stapedio-temporale, new
terms. While arteria facialis is sometimes used as a name for the
stapedial artery of lizards, it has also been used in human anato-
my in a different sense; therefore, Nick's name is avoided here.

The larger passageway from the supraeolumellar portion of
the fenestra postotica leads forward and mesiad to open just
lateral to the notch (incisura pro-otica) in the pro-otic bone for
the trigeminal ganglion. Nick's (1913) felicitous name for this
passageway is canalis cavernosus, while he terms the anterior
orifice the foramen cavernosum : since the chief occupant of the
passageway and the foramen is that portion of the vena capitis

26 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

lateralis homologous with the cavernous sinus of human anatomy,
Nick's terminology is here adopted. Essentially, however, the
canalis cavernosus of turtles corresponds to the cranioquadrate
space or passage of other gnathostomes ; the outer wall is formed
mainly by the quadrate (with some contribution from the ptery-
goid, particularly anteriorly), while the bony ear-capsule forms
the inner wall. Thus, the facial nerve also runs through the
canalis cavernosus from a foramen in the mesial wall of the
canalis.

As to the infracolumellar portion of the fenestra postotica,
the ventro-mesial corner of the fenestra is set off by a snag of the
pterygoid as the foramen caroticum internum of Siebenrock
(1897). In the present paper, I use "stapedial artery" for what
Siebenrock termed the external carotid, and so "internum" be-
comes an unnecessary adjective. It seems desirable, however, to
make a distinction between a foramen into the pituitary fossa for
the carotid, a constant feature of all turtles, and this outermost
opening in the skull for the carotid, an opening that shows in-
teresting and taxonomically useful variations. I here propose the
name foramen caroticum definitivum for the outermost opening
in the skull for the carotid (that is, the internal carotid of many
authors) ; for the foramen into the pituitary fossa (that is, in
what ossifies as the basisphenoid) I shall use the name foramen
caroticum primitivum (this terminology by analogy to that used
for the foramen stylomastoideum in mammals).

The foramen caroticum definitivum leads to a canal that runs
forward to the foramen caroticum primitivum. This canal is
here called the canalis caroticus. The ventral wall of the canalis
caroticus is formed by the pterygoid, the dorsal wall by the ear-
capsule ; that is, the pro-otic forms the dorsal wall anteriorly, and
in Batagur, Callagur, and Kachuga trivittata the opisthotic forms
a more posterior continuation of the dorsal wall (and hence, in
these genera, the dorsal margin of the foramen caroticum definiti-
vum). In Morenia and Geoclemys the pterygoid comes to sur-
round the canalis caroticus, and the foramen caroticum
definitivum is entirely in the pterygoid. The canalis caroticus
gives off two small branches : a posterior branch is given off about
opposite the fenestra ovalis to run forward in the pterygoid bone,
usually to open onto the dorsal surface of the palate near the an-
terior extremity of the descending process of the parietal or just
anterior to the epipterygoid bone; an anterior, usually minute

MCDOWELL: ARTERIES OF TESTUDINOID SKULLS 27

branch issues from the canalis earoticus just lateral to the fora-
men caroticum primitivum or forward of that point and runs
forward on the dorsal surface of the pterygoid to disappear or
else connect by a diagonal groove with the anterior opening of
the canal for the more posterior branch. The more posterior of
these canals is the Vidian canal, while the more anterior is here
called the canalis arteriopalatinus.

In addition, near the foramen caroticum primitivum there may
be a ventral branch of the canalis earoticus that opens onto the
pharyngeal surface of the pterygoid. This is here called the can-
alis carotico-pharyngealis, and its ventral opening, the foramen
carotico-pharyngeale. This is a conspicuous foramen in Ameri-
can Clemmys, Terrapene, Emys, and Emydoidea, but is small or
absent in the other genera.

Arteries : The arteries in the head of Emys orbicularis and
Testudo (Testudo) graeca have been described by Shindo (1914)
and his findings are summarized in Hafferl (1933). Unfortun-
ately, the few papers on head arteries of turtles have been
morphological studies aimed at discovering the basic agreement
between all reptiles, and at interpretation of the fate, in the
adult, of the aortic arches. While these works have been invalu-
able in understanding the broad sweep of anatomical specializa-
tion and evolution among tetrapods, they are less helpful in
understanding the details of phylogeny within one superfamily of
testudinoids.

My most satisfactory dissections have been of two beautifully
injected Chrysemys (Trachemys) scripta prepared by Champlain
Biological Supply Company, to whom I bear a great debt of
gratitude. Less satisfactory dissections (but adequate at least for
the major arteries) have also been made of Chelydra (juvenile),
Rhinoclemmys pulcherrima, Ocadia, Chinemys, Testudo, Gopher -
us, Chrysemys picta, and Malaclemys (Graptemys) geographica.

After giving off the lingual artery (external carotid of some
authors, or ventral carotid) in the anterior part of the neck, the
carotid stem continues forward to divide into two branches, a
dorsolateral stapedial artery and a ventromedial carotid (or
internal carotid). Of the two, the carotid is the smaller.

Almost immediately the stapedial artery gives off small
branches to the region of the Eustachian tube, particularly to a
structure characteristic of all turtles, but never, to my knowledge,

28 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

named or described: the operculum tubae, new term. The oper-
culum tubae is a dense, crescentic pad of connective tissue
attached by its straight dorsal edge to the posterior extremity of
the pterygoid and a roughened area of the quadrate just above
the notch for the columella. The convexly rounded ventral bor-
der of the operculum tubae hangs down as a free flap held only
by loose connective tissue. This flap lies immediately behind the
chorda tympani and the Eustachian tube (it is, in all likelihood,
a specialization of tissue associated with the posterior wall of the
Eustachian tube). The effect of this pad of connective tissue is
to press the Eustachian tube closed ; however, a branch of the
depressor mandibulae muscle (here called the musculus dilator
tubae, new term) runs from a crest formed by the opisthotic and
supratemporal (squamosal of most authors) to the posterior
surface of the operculum tubae. Contraction of the muscle draws
back the operculum and opens the Eustachian tube. Very likely
this is a necessary means of opening the Eustachian tube in a
skull with an immovable pterygoid bone, where kinesis of the
pterygoid will no longer open and close the adjacent pharyngeal
opening of the Eustachian tube. Mammals, faced with a similar
problem, have solved it in a different way : by enwrapping the
Eustachian tube in the swallowing muscles of the velum pala-
tinum, they have made the act of swallowing simultaneously the
act of opening the Eustachian tube by muscular massage.

The stapedial artery (arteria temporomaxillaris of Shindo)
enters the fenestra postotica in a dorsolateral position, near the
dorsal corner of the fenestra. Here, within the skull, between
the quadrate and the bony auditory capsule, the artery divides
into a dorsal and a ventral branch.

The ventral (mandibular) branch runs with the vena capitis
lateralis to exit at the trigeminal aperture. It is usually minute,
but large in Testudo, Gopherus, and Rhino clemmys.

The dorsal (larger) branch passes through the canalis stape-
diarterialis and out by the foramen stapedio-temporale into the
temporal fossa. It is accompanied by a small branch of the vena
capitis lateralis. At the foramen stapedio-temporale the artery
is divided into a posterior branch running back to the neck along
the floor of the temporal fossa (arteria cervicalis of Shindo), and
an anterior branch that curves upward and forward and lies in
a distinct groove on the external surface of the descending process
of the parietal. At about the transverse level of the foramen for

MCDOWELL: ARTERIES OF TESTUDINOID SKULLS 29

the fifth nerve (but considerably dorsal to the nerve foramen)
this anterior branch becomes separated from the parietal and
runs through the musculature of the temporal fossa to a point
dorso-posterior to the eye ; here the artery gives off branches. A
small branch (supraorbital) runs above the eye to supply the
upper lid and to disappear into a small foramen in the prefrontal,
but the main branches (infraorbital) run downward to make con-
tact with the branches of the trigeminal nerve. It is these
branches that supply the palate and nasal chamber, essentially
following the branches of the nerves. The palate is supplied by
a U-shaped artery, continuous across the midline, that runs medial
to the border of the triturating surface. This U-shaped artery
receives blood mainly from a branch of the suborbital artery that
passes through the suborbital foramen or fenestra1 while a branch
of the suborbital artery may join the U shaped artery anteriorly
by passing through the nasopalatine foramen. On each side of
the mid-line the U-shaped artery gives off a forward branch that
enters the nasal chamber from beneath by passing through a pair
of foramina (usually near the premaxillo-vomerine suture) that
have been called the incisive foramina, but since the incisive
foramen of mammalian anatomy is for the duct or ducts of the
nasal gland and Jacobson's organ (which are not associated with
the foramen of turtles), the foramen cannot be homologous in
turtles and mammals. I prefer Seydel's name foramen praepala-
tinum for the foramen of turtles.

The carotid proper (internal carotid) enters the canalis caro-
ticus by the foramen caroticum definitivum, in company with the
auricular and Jacobson's nerves.2 These nerves enter the Vidian
canal to join with the palatine ramus of the facial nerve, forming
the Vidian nerve ; a branch of the carotid, here called the Vidian
branch, also enters the Vidian canal to pass forward to the dorsal
surface of the palate. The Vidian branch of the carotid is very
small. Farther forward, the carotid may give off a small branch
that passes through the canalis carotico-pharyngealis to supply
the roof of the pharynx. Just lateral to the foramen caroticum
primitivum a usually small artery is given off from the carotid to

il term this hole "suborbital foramen" when it is completely filled by the
vessels and nerves as in Malaclemys. When the hole in the dried skull is en-
larged and mainly tilled by membrane in life (e.g. Clemmys) I term it the
"suborbital fenestra."

2i prefer the name "Jacobson's nerve" to "tympanic nerve" for this part of
the sympathetic trunk connecting the glossopharyngeal and palatine nerves, since
in turtles this nerve is remote from the tympanic cavity.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

AOV+SF

Figure 1. Dorsal view of arterial circulation (left side) and arterial
foramina (right side) : testudinid pattern (semidiagrammatic), based on
Malaclcmys (Graptemys) gcographica. Abbreviations: AOV+SF, anterior
orifice of Vidian canal and suborbital foramen; APF, foramen arterio-
palatinum; CA, cerebral carotid artery; FC, foramen cavernosum; IA,
infraorbital artery; ICA, (internal) carotid artery; MA, mandibular artery;
MAF, maxillary arterial (alveolar?) foramen; PA, palatine artery; PMA,
pseudomandibular branch of infraorbital artery; SA, supraorbital artery;
STA, stapedial artery; TA, temporal portion of stapedial artery; TEA,
trigeminal aperture; VA, Vidian artery.

MCDOWELL: ARTERIES OF TESTUDINOID SKULLS 31

run forward in the canalis arteriopalatinus of the pterygoid
(usually a tiny groove, rather than a closed canal). To judge
from the size of the bony foramen, this is a moderately large
artery in Batagur ,C allagur , Kachuga, Morenia, Har delta, Orlitia
and Siebenrockiella. This artery has the position of the artery in-
dicated by grooves on the parasphenoid of labyrinthodonts and
there called the palatine artery,' and accordingly, I term it the
palatine artery here, although in most Testudinidae it appears to
be a vestigial, almost functionless structure. The artery of much
larger size seen in sea turtles and called the arteria palatino-
nasalis by Nick (1913) appears to be homologous in that it is a
forwardly directed branch of the carotid that is separated from
the cerebral carotid by the trabecula cranii (represented in the
adult by the lateral border of the pituitary fossa of the basis-
phenoid). There is usually an anastomotic artery between the
Vidian and palatine arteries.

The carotid enters the foramen caroticum primitivum to pass
into the pituitary fossa. Here there is a small anastomosis be-
tween the left and right arteries behind the pituitary (presum-
ably, a circle of Willis), but the main artery (now the arteria
carotis cerebralis) breaks up into branches in the meninges.

To generalize, apart from vestigial twigs, the separation be-
tween carotid and stapedial arteries in Testudinidae and
Chelydra represents a segregation of blood for the brain (carotid)
from blood for the rest of the head (stapedial artery). Further-
more, the stem of the supraorbital and infraorbital arteries is
extracranial.

II. THE PATTERN OF THE KINOSTERNIDAE

Bony canals and foramina : — I have been able to examine
skulls of Kinosternon, Sternotherus, Claudius, Staurotypus, and
Dermatemys. The most conspicuous difference from the testu-
dinid condition is the absence of the canalis stapediarterialis and
foramen stapedio-temporale, or else the reduction of the canal to
a very small pore. The foramen caroticum definitivum is relative-
ly larger than that of testudinids, as is the canalis caroticus. The
canalis caroticus leads forward to the level of the pituitary fossa
and there divides into two openings : the outer and larger opening

'For a brief review of the position of the groove for the palatine (= palatal =
nasopalatine) artery in labyrinthodonts see Romer (1947, pp. 44-45, fig. 8).

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

RPA

APF

FCD

Figure 2. Dorsal view of arterial circulation (left side) and arterial
foramina (right side) : kinosternid pattern (semidiagrammatic), based on
Kinosternon sonoriense. Abbreviations as in Figure 1 ; also, AOV, anterior
opening of Vidian canal; FCD, foramen caroticum definitivum; EPA,
recurrent pharyngeal branch of Vidian artery (a small artery supplying
pharyngeal epithelium and passing through the pterygoid by the foramen
carotico-pharyngeale) ; SF, suborbital foramen.

MCDOWELL: ARTERIES OF TESTUDINOID SKULLS 33

is that of the canalis arteriopalatinus, while the smaller, inner
opening is the foramen caroticum primitivum. The opening of
the canalis arteriopalatinus is ventral to the foramen cavernosum
and is separated from that foramen by an horizontal lamina of
the pterygoid. For convenience Ave may call this anterior opening
of the canal the foramen arteriopalatinum, new term.

The Vidian canal is small and runs forward from an opening
in the floor of the canalis arteriopalatinus. The Vidian canal is
entirely buried in the pterygoid bone.

Arteries. I have dissected an injected Sternotherus odoratus,
two uninfected Kinosternon subrubrwm, and one uninjected
Kinosternon sonoriense. The stapedial artery appears to be all but
absent ; in the Sternotherus and K. sonoriense a minute artery ex-
tended dorsally from the carotid stem behind the head and
supplied the muscles at the rear of the temporal fossa ; this prob-
ably represented the stapedial. In the Kinosternon specimens,
branches of the vena capitis lateralis passed from the temporal
fossa into a very small vestige of the foramen stapedio-temporale.
Probably the occasional vestiges of this foramen seen in other
Kinosternidae are similarly purely venous.

The carotid divides into a smaller arteria carotis cerebralis
passing through the foramen caroticum primitivum and similar
to that of testuclinids, and a larger arteria palatina passing
through the foramen arteriopalatinum. The blood for head struc-
tures other than the brain comes from the palatine artery. The
palatine artery gives off a main (mandibular) branch that
accompanies the mandibular nerve and supplies the jaw muscles,
then, still within the cavum epiptericum, the palatine artery con-
tinues forward as a large vessel to the orbital region, where it
breaks up into supraorbital and infraorbital branches distributed
as in Testudinidae.

To generalize, in Kinosternidae nearly all the blood of the head
passes through the canalis caroticus, and it is not until the
single main head artery reaches the trabecula cranii and foramen
caroticum primitivum that there is a segregation of blood for the
brain from blood for the remainder of the head. This segregation
takes place not behind the skull, as in Testudinidae, but within
the cavum epiptericum. Further, the stem of the infraorbital
and supraorbital arteries is intracranial.

34 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

III. DISCUSSION

In terms of the bony canals for arteries, the Testudinoidea
(sense of Romer 1956) fall into two groups. One group contains
Chelydra, Macroclemys, and the Testudinidae of Romer (1956).
The other group contains Dermatemys, the Staurotypinae, and
the Kinosterninae of Romer (1956). In testudinids there is
a large foramen stapedio-temporale, but the foramen arterio-
palatinum is small or minute. In the kinosternines the foramen
stapedio-temporale is minute or absent, while the foramen arterio-
palatinum is very large and conspicuous.

It would seem, since blood for the head other than the brain
must come through one or the other of these foramina (except in
land tortoises and some geoemydides,1 where the mandibular
branch of the stapedial is large), that reduction of the one fora-
men must lead to increase in the other. However, it does not
seem likely that the testudinid condition is derived from the
kinosternine pattern, for a large foramen stapedio-temporale,
and, by inference, a well-developed temporal branch of the
stapedial artery, is found in all turtles except the Dermatemys-
Kinosterninae-Staurotypinae series, even in the Amphichelydia ;
moreover, this, the testudinid, type of stapedial artery is the type
seen in Squamata, Rhynchocephalia, and Crocodilia.

On the other hand, the reduction of the palatine artery and
consequent reduction of the foramen arteriopalatinum seen in
Testudinidae and Chelydrinae is also a departure from the con-
dition in other turtles and indicates a specialization in the oppo-
site direction from that of kinosternines. Even in Morenia, Orlitia,
and Batagur, where the foramen arteriopalatinum is the largest
to occur in testudinids, the foramen is decidedly smaller than the
foramen caroticum primitivum. Both arterial types might be
derived from a pattern such as that of sea turtles (see Nick 1913
and Shindo 1914), where there is both a strong stapedial artery,
primarily supplying the jaw musculature, and a strong palatine
artery, primarily supplying the nasal and palatal region. The
kinosternine pattern may be derived from this, probably primi-
tive, pattern, by capture of the temporal circulation by the
palatine artery, while the testudinid pattern may be derived by
capture of the nasal and palatal circulation by the stapedial
artery.

il use the ending "-ide" for a suprageneric taxon to which I cannot yet assign
with confidence a place in the taxonofnic hierarchy.

MC DOWELL : ARTERIES OP TESTUDINOID SKULLS 35

Within each of the two groups of Testudinoidea, as defined by
head arterial foramen pattern, there is relatively little variation.
The most notable is that seen in Gopherus and some species of
Gcochclone (e.g., G. partialis and G. denticulata, but not G. gi-
gantea and G. radiata) . Here, the lower (mandibular) ramus of
the stapedial artery does not accompany the vena capitis lateralis
anteriorly, but makes exit to the temporal fossa through its own
foramen, here called foramen arteriomandibularis, new term,
lying lateral to the foramen cavernosum. Other than this, the
land tortoises are very similar to pond turtles, and their classifi-
cation together in one family, Testudinidae, seems thoroughly
warranted. (A foramen arteriomandibularis occurs as a varia-
tion in Bhinoclemmys and is always potential where the ramus
mandibularis is large. )

The present findings do not, however, support the classification
of Williams (1950) or its expanded form given in Homer (1956)
in all details. This classification placed Dermatemys in a separ-
ate family (along with certain poorly known fossil genera), the
Dermatemydidae ; Chelydra and Macroclemys (syn. Macrochclys)
formed the Chelydrinae, Claudius and Staurotypus the Stauro-
typinae, and Kinosternon and Sternotherus the Kinosterninae of
a family Chelydridae;1 Platysternon formed the Platysterninae,
the pond turtles the Emydinae, and the land tortoises the Testu-
dininae of a family Testudinidae. While the foramina for arteries
in the main conform to this classification, particularly on such
points as the close relationship between Platysternon, the pond
turtles, and the land tortoises, and the close relationship between
the Kinosterninae and the Staurotypinae, the arterial foramina
suggest that Dermatemys is much more closely related to the
Staurotypinae than is indicated by Williams' classification, and
that Chelydra and Macroclemys are more closely related to the
Testudinidae than to the remainder of Williams' Chelydridae. As
here interpreted, Chelydra and Macroclemys are already com-
mitted to the testudinid evolutionary line in the reduction of the
palatal artery (much more reduced than in many Old World
emydines), while Dermatemys is committed to the kinosternid
evolutionary line in its loss of the stapedial artery as indicated by
the absence of a foramen stapedio-temporale.

Dermatemys, Staurotypus, and Claudius were grouped to-
gether by Boulenger (1889) in the Dermatemydidae. While later

iThis association of kinosternines, staurotypines., and chelydrines in one family
appears to date from Gray 1870.

36 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

work has demonstrated Boulenger's error in separating Stauro-
typus and Claudius from the kinosternines, the affinity between
Dermatemys and the Staurotypinae suggested by Bonlenger
seems much more real to me than does the suggested affinity of the
staurotypines and kinosternines with Chelydra and Macroclemys.

Several other points suggest that kinosternines are close to
Dermatemys, while Chelydra and Macroclemys are close to the
Testudinidae. Thus the kinosternines and Dermatemys have a
conical and radially symmetrical footplate of the columella auris,
while the footplate in chelydrines is flattened and disc-like, with
the shaft arising from below the center; most testuclinids have
a columella as in chelydrines, but some Asiatic forms (e.g. Mor-
enia, Batagur, Orlitia) approach kinosternines in the conical
form of the footplate, although the asymmetry is marked.

Again, in kinosternines and Dermatemys there is a tight suture
between basioccipital and opisthotic behind the acoustic papilla of
the inner ear ; Chelydra and Macroclemys agree with Testudin-
idae in the presence of an open fissure in the suture behind the
apex of the papilla. Further, cloacal bursae are absent in Der-
matemys and kinosternines, but present in Chelydra and Macro-
clemys; most aquatic Testudinidae have bursae (Smith and
James ) .

The genus Baptemys, of the North American Eocene, appears
to be intermediate morphologically between Dermatemys and the
kinosternines. The genus is known from a number of shells, a few
fragments of the limb skeleton, and two partial skulls. Of the
skulls, one (in the Yale University Peabody Museum) is nearly
complete. I have examined this skull and find it quite as figured
by Hay (1908, pi. 37).

Baptemys agrees with both Dermatemys and the kinosternines
in the absence of the foramen stapedio-temporale, the great
forward extent of the posterior temporal emargination, and the
entrance of the squamosal (quadrato-jugal Auct.) into the border
of the posterior temporal emargination so that the postorbital is
broadly excluded from the supratemporal (squamosal Auct.).
Baptemys agrees with Dermatemys and differs from the kinoster-
nines in: 1) having a moderately deep inferior temporal emar-
gination; 2) having the trochlear process of the crista prae-
temporalis concealed from lateral view by the temporal arch ;
3) having rather large frontals, with the interfrontal suture
longer than the interprefrontal suture. Baptemys agrees with

MCDOWELL: ARTERIES OF TESTUDINOID SKULLS 37

kinosternines and differs from Dermatemys in: 1) absence of
keels on the triturating surfaces; 2) high, acute coronoid process
of the lower jaw; 3) lack of serrations on the jaw margins; and
4) reduction of the plastron. In one feature, Baptemys differs
from both Dermatemys and the kinosternines : there is a complete
series of neurals, separating the pleurals on the midline through-
out the carapace.

It should be noted that the genera among the testudinid group
that come nearest to the Dermatemys-ldnostermne series are not
Chclyelra and Macroclemys, but the Asiatic Batagur -M or enia-
series, where the foramen arteriopalatinum is less reduced than
in other Testudinidae and the footplate of the columella auris is
conical. The general appearance of the skull of Dermatemys is
strongly reminiscent of Batagur and Hardella, particularly in
such features as upturned nares, broad and complexly ridged
triturating surfaces and broad muscular tuberosities of the basi-
occipital. This resemblance extends to the pes, for Dermatemys
agrees with Hardella, Kaehuga, Callagur, and Batagur in having
four phalanges in the fifth toe (usually there are three or two
phalanges in this toe in Testudinoidea) .

Thus Asiatic "batagurs," on the one hand, and the Central
American Dermatemys, on the other, appear to be near the base
of the divergence of the testudinid and kinosternid series.

ACKNOWLEDGEMENTS

For the loan of material, I am grateful to Dr. Doris Cochran
(United States National Museum), Dr. Norman Hartweg (Uni-
versity of Michigan Museum of Zoology), and Dr. Ernest
Williams (Museum of Comparative Zoology). The Champlain
Biological Supply Company most generously provided dissection
material of Chrysemys (Trachcmys) scripta. For aid, both
athletic and moral, in the examination of the Yale Peabody Mu-
seum skull of Baptemys, I am in debt to Mr. Peter Robinson. I
have profited much from discussions of turtle morphology with
Dr. Williams and with Dr. Thomas Parsons. Without the aid of
my wife, Rosa McDowell, in the preparation of the manuscript,
this paper would have been impossible.

38 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

LITEEATUEE CITED

Baub, G.

1888a. Osteologische Notizen iiber Eeptilien (Fortsetzung IV.). Zool.

Anz., vol. 11, pp. 592-597.
1888b. Osteologische Notizen iiber Eeptilien (Fortsetzung V.). Ibid.,

pp. 736-740.

BlENZ, A.

1895. Dermatemys mavii Gray eine osteologische Studie. Eev. Suisse
Zool., vol. 3, pp. 61-136, pis. 2, 3.

BOULENGER, G. A.

1889. Catalogue of the chelonians, rhynehocephalians, and crocodiles
in the British Museum (Natural History). London: British
Museum, x + 311 pp., 73 figs., 6 pis.
Gray, J. E.

1870. Supplement to the catalogue of shield reptiles in the collection
of the British Museum. Part I. Testudinata (tortoises). Lon-
don, 120 pp., 40 figs.
Hafferl, Anton

1933. Das Arteriensystem. In Handbuch der vergleichenden Anatomie
der Wirbeltiere, L. Bolk, E. Goppert, E. Kallius, and W.
Lubosch, eds. Berlin and Vienna: Urban & Schwarzenberg,
vol. 6 (xii + 854 pp., 629 figs.), pp. 563-684, figs. 414-521.
Hay, 0. P.

1908. The fossil turtles of North America. Carnegie Inst. Washington,
Publ. 75, pp. i-iv, 1-568, figs. 1-704, pis. 1-112.

KUNKEL, B. W.

1912. The development of the skull of Emys lutaria. Jour. Morph.,
vol. 23, pp. 693-780.

Nick, L.

1913. Das Kopfskelet von Dermochelys coriacea L. Zool. Jahrb.
(Abt. Anatomie und Ontogenie), vol. 33, pp. 1-238, figs. A-Q,
pis. 1-12.

Eomer, A. S.

1947. Eeview of the Labyrinthodontia. Bull. Mus. Comp. Zool., vol.

99, pp. 1-368, figs. 1-48.
1956. Osteology of the reptiles. Chicago : University of Chicago Press,

xxi + 772 pp., 248 figs.

Shindo, T.

1914. Zur vergleichenden Anatomie der arteriellen Kopfgefasse der
Eeptilien. Anat. Hefte, vol. 51, pp. 267-356, figs. 1-21, pi. 9.

Siebenrock, F.

1897. Das Kopfskelet der Schildkroten. Sitzungsb. Math.-Naturw.
Classe K. Akad. Wiss. Weill, vol. 106, pt. 1, pp. 245-328, pis. 1-6.

MCDOWELL: ARTERIES OF TESTUDTNOID SKULLS 39

Smith, H. M. and L. F. James

1958. The taxonomic significance of cloacal bursae in turtles. Trans.
Kansas Acad. Sci., vol. 61, no. 1, pp. 86-96.
Williams, E. E.

1950. Variation and selection in the cervical central articulations of
living turtles. Bull. Araer. Mus. Nat. Hist., vol. 94, pp. 505-561,
figs. 1-20, tables 1-10.
1952. A staurotypine skull from the Oligocene of South Dakota (Test
udinata, Chelydridae). Breviora, no. 2, pp. 1-16, pis. 1-2.

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Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE

Vol. 125, No. 3

TWO JURASSIC TURTLE SKULLS
A MORPHOLOGICAL STUDY

THOMAS S. PAESONS
Department of Zoology
University of Toronto

And

EENEST E. WILLIAMS
Museum of Comparative Zoology

With Six Plates

CAMBEIDGE, MASS., U.S.A.
PEINTED FOE THE MUSEUM

June, 1961

No. 3 — Two Jurassic Turtle Skulls: A Morphological Study
By Thomas S. Parsons and Ernest E. Williams

Introduction

We here describe the detailed morphology of the skulls of two
Jurassic turtles, one from the Portland Limestone of England, repre-
sented by three partial crania and one mandible (R2914, R3163, and
R3164, British Museum, Natural History), and another from the Solen-
hofen of Bavaria, a cranium and mandible (4023) belonging to the Teyler
Museum, Haarlem, Holland.

We deal with these specimens as examples of the morphology of
turtle skulls in the Upper Jurassic. We do not attempt to assign them
names or to place them taxonomically. The British Museum specimens
have some similarity to Stegochelys planiceps (Owen) which is also from
the Portland Limestone. This may well be what the Portland specimens
should be called, but Stegochelys planiceps (Owen) is a genus and species
founded upon a unique skull, and this type specimen is not at present
to be found; the available figures (Owen, 1849-1884, plate 8) are some-
what idealized and do not compare in detail with the specimens at hand.

In the case of the Solenhofen skull we do not have any hint as to
its proper name. It is surely a very different genus from the Portland
one, exhibiting a remarkable combination of very specialized and primi-
tive characters in contrast to the mostly primitive features of the Portland
species. It is very probably a member of a different family from that to
which the Portland form will be found to belong.

We might very safely provide a new generic and specific name for
the Solenhofen skull, or with still greater security refer the Portland
specimens provisionally to Stegochelys planiceps. We see, however, no
great advantage in so doing. At present the higher taxonomy of Jurassic
turtles is based exclusively upon shells. It would be necessary to have
shells associated with these skulls in order to place them as to family.
To devise or to use skull genera for these forms, when these skull genera
will most probably in the not distant future be sunk in the synonymy of
shell genera, would, we believe, serve no useful purpose.

Even at the familial level, exact assignment is impossible. We would
suggest the following : they are surely amphichelydians and presumably
pleurosternoids. Of the three families at present recognized in the Upper
Jurassic of Europe, the Pleurosternidae, Plesiochelyidae, and Thalas-
semydidae, the first family is the most primitive, while the latter two
are rather highly specialized and closely related ; it would therefore be
plausible to suppose that the Portland form is a pleurosternid and the

44 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Solenhofen form either a plesiochelyid or a thalassemydid. This is, how-
ever, no more than a guess that the more primitive skull will be
associated with the more primitive shell and the more specialized skull
with the more specialized shell.

It should be pointed out that excellent associated skull and shell
material of Jurassic age does exist in European collections. They are,
however, at the moment almost unprepared and convey an absolute
minimum of information. The skulls we here describe were all, when
first seen, embedded in limestone to such an extent that only a turtle
skull or mandible was apparent. Now, after preparation by acetic acid,
even small cranial foramina may be probed with bristles and only the
erosion of certain parts before fossilization prevents the description
of these specimens from being as complete as that of any Recent skull.
Many untouched skulls in pure limestones of Jurassic age would yield
quite as well to similar techniques. Substantial advances in our knowledge
await only the utilization of present opportunities.

Morphology

A detailed study of the structure of these specimens is presented
in the following pages. We have found it convenient in places to contrast
the Portland and Solenhofen materials by the device of parallel columns.
In all such cases the Portland specimens are discussed on the left and the
Solenhofen on the right.

The Skull as a Whole

In none of the available specimens is the skull roof well preserved.
Posterolaterally, it is partially or completely lost, and the remainder
is often somewhat cracked, thus making a detailed description of the
general shape impossible. The dorsal surfaces of the roofing bones are
rather rugose, though without well-defined sculpture, in the Portland
skulls; they are quite smooth in the Solenhofen skull. However, in
neither case can the sutures be found Avith any degree of certainty. In
both forms, the skull is moderately high in lateral view and triangular
when seen dorsally. The Solenhofen skull is somewhat lower and wider.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

PORTLAND SPECIMENS

The roof of the fossa temporalis
is mostly lost in the specimens, but
was probably fairly complete. At
least there is no indication of any
marked emargination ; however, the
posterior and ventrolateral margins
are not present. The orbits are
quite large and face nearly directly
laterally. Since their margins are
incomplete, the shape cannot be
described.

Most of the dorsal surface of the
skull is nearly horizontal, sloping
only slightly anteriorly. However,
there is a very distinct angle in the
region of the posterior end of the
nasals and the anterior portion of
the face faces anterodorsally, lying
at approximately a forty-five de-
gree angle to the horizontal. Thus
the single median apertura narium
externa faces anterodorsally. When
seen in dorsal or ventral view, the
part of the skull anterior to the
orbits forms an equilateral triangle.

The primary palate possesses
rather narrow, but much roughened
triturating surfaces which are sepa-
rated by a deep median trough into
which the aperturae narium in-
ternae open. Each pterygoid pos-
sesses a well developed processus
pterygoideus extemus; thus the
fenestra subtemporalis is compara-
tively small and posteriorly located.
The processus articularis of the
quadrate is well developed, and
projects well ventral to the surface
of the palate. It lies anterior to the
level of the condylus occipitalis.

SOLENHOFEN SPECIMEN

The roof of the fossa temporalis
is at least moderately complete. It
is lost posteromedially and ventro-
lateral^, and was probably some-
what emarginate in the latter area ;
the posterior margin of the maxilla
does not appear to be broken. The
circular orbits are of moderate size
and face dorsolaterally.

Most of the dorsal surface of the
skull slopes anteriorly, at an angle
of approximately thirty degrees,
and there is no sharp angle when
the skull is seen in lateral view.
Thus the single median apertura
narium externa faces more dorsally
than anteriorly. The anterolateral
surface of the skull is quite concave
when seen in dorsal or ventral
view; the snout, therefore, appears
to be rather long and narrow.

There is a rather extensive and
flat secondary palate formed large-
ly by the maxillae, but also entered
by the palatines. The posterior
end of this palatal surface is
notched, and forms the anterior
margin of an oval median trough
into which a single apertura nari-
um interna opens. There is almost
no trace of a processus pterygoideus
extemus on the pterygoid. Thus
the fenestra subtemporalis is very
large, reaching from the posterior
margin of the maxilla to the proces-
sus articularis of the quadrate. The
latter process is very short so that
the wide condylus mandibularis is
at essentially the same level as most
of the palate ; it projects somewhat
posteriorly so that the mandibular
and occipital condyles all lie in a
straight transverse line.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

FCI"J CAJ

Fig. 1. Portland skull, British Museum E2914. Palatal view. Abbreviations:
CAJ, cavum acustico-jugulare. FCI, foramen posterior eanalis carotici interni. FON,
foramen orbito-nasale. FPP, foramen palatinum posterius. FPR, foramen praepalatin-
um, PPE, processus pterygoideus externus. PTE, processus trochlearis. V, vomer.

Posteriorly, in each the cavum tympani is well developed, and much
like that of recent turtles. It is connected to the large fenestra postotica
by an incisura columellae auris. The massive paroccipital extension of the
otic capsule characteristic of turtles is also like that of recent forms. The
crista supraoccipitalis is broken off so that its extent and shape cannot be
determined.

Skull Roof
Os parietale

The parietals roof over the median portion of the posterior half of
the skull. Anterodorsally, they meet the frontals in a transverse suture
near the midline, and, farther laterally, are in contact with the post-
orbitals. Their posterior and posterolateral portions are either missing

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

— V FPR

Fig. 2. Solenhofen skull, Teyler Museum. Palatal view. Abbreviations: BS,
basisphenoid. FCI, foramen posterior canalis carotici interni. FPP, foramen
palatinum posterius. FPE, foramen praepalatinum. PAL, palatine. PPE, processus
pterygoideus externus.

or have no visible sutures. In neither form is there any extensive posterior
emargination of the parietals; whether or not a small one was present
cannot be determined. Paramedially the parietal sends down a vertical
processus inferior parietalis which meets the supraoccipital posteriorly,
the prootic, and the epipterygoid anteriorly. This process forms the
dorsal wall of the large foramen nervi trigemini.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The parietal is in contact with The parietal is completely sepa-

the crista pterygoidea only at the rated from the pterygoid by the
anterior and posterior margins of well-developed epipterygoid anteri-
the foramen nervi trigemini ; an- orly and by the prootic posteriorly,
teriorly the parietal and the ptery-
goid are separated by the large
epipterygoid.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Os frontale

PORTLAND SPECIMENS

The frontal appears (the sutures
are very obscure) to be a small
trapezoidal element when seen in
dorsal view. It is bounded post-
eriorly by the parietal, medially by
the other frontal, anterolateral^
by the prefrontal, and laterally by
the postorbital. It does not, if this
interpretation is correct, enter the
margin of the orbit.

SOLENHOFEN SPECIMEN

The sutures are very indistinct,
and the shape of the frontal as seen
dorsally cannot be described with
certainty. It appears to be bounded
by the parietal posteriorly, by the
other frontal medially, and by the
postorbital posterolaterally. It
probably does enter the margin of
orbit. Anteriorly the bone is too
eroded for any sutures to be visible.

Each frontal possesses a small parasagittal ridge along its ventral
surface ; between these ridges is a narrow sulcus olfactorius which is
widely open ventrally.

SOLENHOFEN SPECIMEN

Due to the erosion of the antero-
dorsal part of the skull roof, the
shape of the horizontal plate of the
prefrontal cannot be determined.

Os pracfrontale

The prefrontal consists of a horizontal plate on the dorsal surface of
the skull and a large, more or less vertical plate extending ventrally to
meet the bones of the palate. The latter plate forms the bony septum
between the orbit and the fossa nasalis.

PORTLAND SPECIMENS

In dorsal view the prefrontal is
a rather irregularly shaped bone,
the boundaries of which are not al-
ways clear. The horizontal plate is
very thick. Medially it is bounded
by the other prefrontal, postero-
medially by the frontal, postero-
lateral^7 by the postorbital, lateral-
ly by the orbit, anterolateral!}7 by
the maxilla, and anteromedially by
the nasal. These sutures appear to
be essentially straight except for
that with the maxilla which runs
posteriorly from the nasal-pref ront-
al suture and then turns abruptly
anterolateral^ to run to the an-
terior end of the orbit. From there
it extends ventrally between the
vertical plate of the prefrontal and
the maxilla.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

Along the ventral surface of the prefrontal the sulcus olfactorius
continues anteriorly from the frontal.

The vertical plate forms a broad, roughly transverse sheet of bone,
the ventral end of which curves posteriorly, especially in its medial half.
Laterally, the entire length of the plate possesses a suture with the
vertical portion of the maxilla ; medially, it forms the dorsal three-
quarters of the margin of the fissura cthmoidalis. This fissure, which is
roofed by the horizontal plates of the prefrontals, is quite wide dorsally
and becomes gradually narrower ventrally. The posteroventral margin
of the vertical plate is somewhat notched by the anterior margin of the
foramen orbito-nasale.

PORTLAND SPECIMENS

Medial to this foramen, the pre-
frontal is broadly joined with a
small dorsal process on the vomer
anteromedially and, farther pos-
terolateral^, with the anteromedial
margin of the palatine. The fora-
men orbito-nasale is large and
circular.

SOLENHOFEN SPECIMEN

Medial to this foramen, the pre-
frontal forms a suture with a small
dorsal process on the vomer. It
probably also joined the palatine
as in the Portland skulls ; however
the anterior portion of the palatine
is broken in the specimen so that
there is now no contact, and the
shape of the foramen orbito-nasale
cannot be determined.

Os nasale

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The nasals are small, roughly The nasals are not known ; due

rectangular elements forming the to the erosion of this portion of the

roof of the fossa nasalis. Anterior-
ly, they form the essentially trans-
verse dorsal border of the apertura
narium externa. Their sutures are
not clear, but each appears to
possess a straight parasagittal con-
nection with the maxilla laterally,
a sagittal suture with the other
nasal medially, and a roughly
transverse one with the prefrontal
posteriorly. The nasals are thickest
posteriorly, and taper to a thin
edge anteriorly.

skull it cannot even be seen whether
or not they were present.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Os postorbitale

The postorbital is not well preserved in either form. Parts of it
are missing, and the sutures are very indistinct. Thus its posterior and
ventral portions cannot be described.

PORTLAND SPECIMENS

The postorbital probably pos-
sessed an anteromedial suture with
the prefrontal (thus excluding the
frontal from the margin of the
orbit), a medial suture with the
frontal, and a posteromedial one
with the parietal.

SOLENHOFEN SPECIMEN

The postorbital was probably
smaller than in the Portland form,
and was bounded anteriorly by the
frontal (which thus enters the
margin of the orbit).

Os jugale

PORTLAND SPECIMENS

The jugal is not well known since
this area is almost completely lost.
Part of it is preserved in R3164.
and appears to lie between the
maxilla and the postorbital at the
posteroventral corner of the orbit.

SOLENHOFEN SPECIMEN

The sutures are difficult to see,
but the jugal is apparently a rather
small element, although its poste-
rior edge is probably broken so that
its full extent is uncertain. This
bone forms the posteroventral wall
of the orbit, having a vertical
portion which extends ventrally
from the postorbital to the maxilla ;
it is triangular in section. The
jugal apparently lies dorsal to the
maxilla, and does not reach the
outer margin of the triturating sur-
face or the palate (the suture with
the maxilla cannot be seen on the
outer side of the cheek). A fairly
short but stout process runs pos-
teromedially from the vertical por-
tion, bounded ventrally by the
maxilla and medially by the pala-
tine. The end of this process lies
on the dorsal surface of the
processus pterygoideus extemus of
the pterygoid. A small foramen
enters the suture between the jugal
and the palatine near the posterior

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

SOLENHOFEN SPECIMEN

end of this process, and two other
foramina are present well ventrally
on the posterior face of the vertical
portion of the jugal.

Os quadrato- jugate

The quadrato-jugal is not known ; this area is mostly lost, and badly
broken and eroded where present.

Os squamosum

PORTLAND SPECIMENS

The squamosal is not known with
certainty, but is probably repre-
sented by two fragments of bone,
one posterior and one posterodorsal
to the margin of the cavum tympani
of the quadrate. The more poste-
rior of the fragments is in contact
with the extreme lateral end of the
opisthotic posteroventrally. There
is a small hole between the postero-
dorsal part of the cavum tympani
and the posterior end of the more
dorsal of the two fragments ; pre-
sumably this represents the remains
of an antrum postoticum.

SOLENHOFEN SPECIMEN

The squamosal forms a small cap
of bone lying above and behind
the posterodorsal third of the mar-
gin of the cavum tympani of the
quadrate. Anterolateral^, it is
continuous with another (unidenti-
fiable) bone of the skull roof. Its
ventral margin lies on the quadrate
except at the extreme posteromedial
corner where it meets the opisthotic
on the floor of the fossa temporalis.
The posterior extremity of the
squamosal is missing in tbis speci-
men. The bone is everywhere thin
and contains a moderately large
antrum postoticum which is widely
open into the cavum tympani.

Palatal Elements
Os praemaxillare

The premaxilla is a small, roughly triangular element which is
narrowest posteriorly. Medially, it is bounded by the other premaxilla
and posterolateral^ by the maxilla.

PORTLAND SPECIMENS

The anteriormost end is missing
so that it is impossible to know
whether the tip of the upper jaw
was hooked, notched, or plane.

Both the tomial ridge and the

SOLENHOFEN SPECIMEN

The anterior margin of the pre-
maxilla is rather gently curved
when seen in ventral view and it
is neither hooked nor notched.

Lingual ridges are not present,

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS

lingual ridge are large; between
them there is a deep channel on
the palatal surface. The ridges and
channel are continuous with those
of the maxilla. There is, in the
midline, a narrow groove separat-
ing the lingual ridges of opposite
sides. Posteriorly, the palatal sur-
face of the premaxilla ends in a
short transverse suture with the
vomer. This suture is well dorsal
to the level of the lingual ridges.
Lateral and anterolateral to the
vomeral suture, the premaxilla
forms the anterior margin of the
nearly circular foramen praepa-
latinum. At the anterolateral
corner of this foramen, the pre-
maxilla just barely touches the
vomer again, thus excluding the
maxilla from the margin of the
foramen ventrally.

SOLENHOFEN SPECIMEN

and the palatal surface is quite
smooth, although it forms a sharp
tomial margin. The two pre-
maxillae appear as a long wedge,
bounded for their entire postero-
lateral margin by the maxilla; the
vomer does not appear on the
palate. Just inside the tomial
margin on the palatal surface of
each premaxilla, there are three
prominent nutritive foramina. Near
the posterior end of the maxillary -
premaxillary suture there is, on
either side, a very small foramen
leading to the fossa nasalis. They
are not symmetrical and could be
the result of weathering of the
skull, but are more probably very
small vestiges of the foramina prae-
palatina. On one side, the foramen
appears to be completely surround-
ed by the premaxilla, and, on the
other, the maxilla enters its margin.

Dorsally, the premaxillae form the ventral margin of the single
apertura narium externa which is essentially transverse with the lateral
corners slightly rounded. There is no indication of any bony separation
of the two nares extemi. Running along the midline between the anterior
facial surface of the premaxilla and its suture with the vomer dorsally,
there is a markedly thickened and raised portion of the bones; lateral
to this, each is much lower and thinner. At the anterior end, this ridge
is divided by a median (nutritive?) foramen entering the bone of the
snout.

PORTLAND SPECIMENS

To each side of the median ridge
there is a marked groove running
from the foramen praepalatinnm to
a large anterior foramen which con-
tinues anteriorly into the thick
anterior face of the premaxilla.
Besides forming the anterior and

SOLENHOFEN SPECIMEN

The median ridge widens anteri-
orly where it surrounds the median
foramen and overlies a small
groove. This groove appears to lead
into a canal running anteriorly
within the premaxilla. The vomer
meets the premaxilla only at the

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

PORTLAND SPECIMENS

medial margins of the foramen
praepalatinum, the premaxilla
sends a very narrow process along
its lateral margin to meet the vomer
posteriorly and thus exclude the

maxilla from
foramen.

the margin of the

SOLENHOFEN SPECIMEN

top of the ridge ; laterally the
maxilla and premaxilla are in con-
tact. The foramina praepalatina
appear essentially the same as they
do in a palatal view.

Fig. 3. Portland skull, British Museum R2914. Bones within apertura narium
externa (edge broken) as seen in anterior view. Abbreviations: FE, fissura ethmoidalis.
PRF, prefrontal. S V, sulcus vomeri.

PM FPR

Fig. 4. Portland skull, British Museum R2914. Bones within apertura narium
externa (edge broken) as seen in anterodorsal view. Abbreviations: FPR, foramen
praepalatinum. PM, premaxilla. SV, sulcus vomeri. V, vomer.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Os maxillare

The maxilla is a large element with two main portions, a vertical
part and a horizontal part. The former forms the surface of the face
anterior and ventral to the orbit. Anteroventrally, it is bounded by the
premaxilla and, posterodorsal to the latter, it forms the lateral margin of
the apertura narium externa.

PORTLAND SPECIMENS

Posterodorsal to the narial open-
ing it is bounded medially by the
nasals and finally, at its dorsalmost
end, by the prefrontal. The suture
with the prefrontal continues
ventrally along the anterior margin
of the orbit. The maxilla continues
posteriorly as a rather narrow wall
ventral to the large orbit. The
posterior portion is incompletely
preserved, but appears to be
bounded posterodorsally by the re-
mains of the jugal. The lateral
surface of the vertical plate of the
maxilla possesses only a very few
nutritive foramina.

SOLENHOFEN SPECIMEN

The dorsal extension of the
maxilla between the narial opening
and the orbit is eroded and the
sutures cannot be seen. At the
anteroventral corner of the orbit,
it possesses a suture with the pre-
frontal as in the Portland form.
The maxilla continues posteriorly
as a deep bar ventral to the rather
small orbit. It has a suture with
the jugal along the posteroventral
margin of the orbit. Farther
posteriorly, sutures cannot be seen.
There are large numbers of nutri-
tive foramina along the lateral
surface of the maxilla.

Internally, the maxilla forms the lateral and ventrolateral walls of
the fossa nasalis. At the posterolateral corner of the nasal fossa and at
the lateral edge of the foramen orbito-nasale, lies the foramen alveolare
superius which leads into the canalis alveolaris superior ; the various
nutritive foramina of the facial and palatal surfaces of the maxilla are
all connected with this canal.

PORTLAND SPECIMENS

The maxilla forms the floor of
the orbit laterally. Anteriorly it is
in contact with the ventrolateral
portion of the prefrontal and pos-
teriorly with the processus ptery-
goideus externus of the pterygoid.

SOLENHOFEN SPECIMEN

The maxilla forms only a small
part of the floor of the orbit antero-
laterally. It is bounded anteriorly
by the prefrontal, anteromedially
by the margin of the foramen
orbito-nasale, posteromedially by

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

SOLENHOFEN SPECIMEN

the palatine, and posterolaterally
by the jugal.

PORTLAND SPECIMENS

Between these two, it forms a
suture with the palatine. The
maxillary suture with the palatine
is separated from that with the
prefrontal by the foramen orbito-
nasale, and its suture with the
palatine separated from that with
the pterygoid by the foramen
palatinum posterius ; the maxilla
forms the lateral margin of both of
these large foramina.

Posterior and lateral to the premaxilla, most of the palate is formed
by the horizontal portion of the maxilla.

PORTLAND SPECIMENS

This is a rather long, narrow
element with a very prominent
tomial ridge and, along its medial
edge, an equally prominent lingual
ridge ; between the two there is a
deep channel. Both the ridges and
the channel are continuous with
those of the premaxilla. The tomial
ridge is sharp but quite even, while
the lingual ridge has an irregular,
roughened surface. The apertura
narium interna lies medial to the
center of the maxilla which forms
its anterolateral and lateral mar-
gins. Anterior to the apertura, the
maxilla extends slightly medially
and dorsally to meet the vomer.
Anterior to the vomer it forms a
suture with the premaxilla and is
narrowly excluded from the margin

SOLENHOFEN SPECIMEN

This is a large plate since this
form has a well-developed second-
ary palate. The tomial ridge is low
but sharp, and the remainder of its
surface is smooth. It is bounded
anteromedially by the premaxilla,
medially by the other maxilla, and
posteromedially by the palatine
which excludes it from the margin
of the apertura narium interna. At
its extreme posterior end it meets
the reduced processus pterygoideus
externus of the pterygoid. The con-
cave posterolateral margin of the
maxilla borders the fenestra sub-
temporal. Numerous, moderate-
sized nutritive foramina occur
along the tomial ridge and scattered
larger ones are present on the
palatal surface. The former give

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

SOLENHOFEN SPECIMEN

the tomial ridge a somewhat serrate
appearance.

PORTLAND SPECIMENS

of the foramen praepalatinum.
Posterior to the apertura narium
interna, it joins the palatine ante-
rior to the large oval foramen pa-
latinum postering and forms the
lateral margin of that foramen.
The processus ptcrygoideus extern-
us of the pterygoid meets the
maxilla at the posterior end of the
deep channel between the tomial
and lingual ridges. In the channel
are a few moderately large nutri-
tive foramina ; there are fewer and
smaller ones on the tomial ridge.
The posterolateral corner of the
maxilla is not preserved.

Os palatinum,

The palatine is a plate of bone forming the portion of the palate
medial to the posterior half of the maxilla.

PORTLAND SPECIMENS

In palatal view the two palatines
are separated by the vomer with
which they are in contact through-
out their entire length. Each forms
a rather flat plate which is farthest
dorsal at its anteromedial corner
where it forms a short suture with
the ventromedial portion of the
prefrontal ; from there it slopes
posteroventrally. Posteriorly i t
possesses a rather irregular suture
with the pterygoid. Laterally, it
forms the medial margin of the
foramen palatinum posterius and
the posteromedial margin of the
foramen orbit o-nasale. Between
these foramina a massive process
extends ventrolateral^ to meet the
posteromedial edge of the maxilla
where it forms a small posterior

SOLENHOFEN SPECIMEN

Due to the formation of a second-
ary palate, each palatine forms a
much folded plate which is, anteri-
orly, C-shaped in transverse section.
It possesses a rather small, lenticu-
lar, horizontal portion along the
posteromedial margin of the second-
ary palate. This portion is bounded
anterolaterally by the maxilla and
posteromedially it forms the mar-
gin of the apertura narium interna.
Anteriorly the two palatines are in
contact to exclude the maxilla from
the margin of the apertura. Post-
eriorly the palatal surface of the
palatine possesses a short suture
with the pterygoid. The palatine
continues dorsally from its line of
suture with the maxilla and then
arches dorsomedially to meet the

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

PORTLAND SPECIMENS

portion of the prominent lingual
ridge. The medial half of the pala-
tine forms the roof of the apertura
narium interna. The vomer extends
somewhat ventral to the medial por-
tion of the palatine so that there is
a distinct groove between the vomer
and the lateral portion of the pala-
tine.

The dorsal surface of the palatine
forms most of the floor of the orbit ;
its relations there are the same as
on the ventral surface except that
the two palatines meet in a sagittal
suture dorsal to the vomer for the
posterior two-thirds or three-quar-
ters of their length.

SOLENIIOFEN SPECIMEN

other palatine in a sagittal suture
along the roof of a deep, median
trough. Thus the apertura narium
interna is surrounded by the pala-
tines. The vertical and dorsal parts
of the palatine continue a short
distance posterior to the maxilla
and are there bounded laterally by
the pterygoid. On the left side
there is a small oval foramen pala-
tinum posteriits between the pala-
tine and the pterygoid ; on the right
it is divided into two foramina, a
more posterior oval one like that on
the left and a smaller circular one
completely within the palatine just
anterior to the first. There are one
or two small foramina near the
posterior end of the secondary
palatal surface of the palatine.
Posteriorly the palatine was almost
certainly in contact with the ptery-
goid, but on the specimen the
posterodorsal portion of the former

is missing.

The dorsal surface of the palatine
forms the ventromedial wall of the
orbit. Anteriorly the dorsal part is
somewhat broken, but meets the
vomer anteromedially. Lateral to
that it almost certainly possessed a
suture with the ventromedial edge
of the prefrontal and formed the
posteromedial margin of the fora-
men orbit o-nasale.

Vomer

PORTLAND SPECIMENS SOLENIIOFEN SPECIMEN

The vomer extends from the pre- The vomer is much reduced and

maxillae to the pterygoids, forming does not appear on the palate at
a long narrow bar along the center all, but forms a short, rather stout

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

SOLENHOFEN SPECIMEN

vertical element lying dorsal to the
sagittal suture between the max-
illae and palatines on the secondary
palate. Thus the vomer separates
the anterior halves of the two meati
choanarum which lead from the
fossa nasalis to the aperturae
narium internae.

PORTLAND SPECIMENS

of the palate. Anteriorly it is a
rather stout element which has a
short transverse suture with the
premaxilla and which forms the
posterior and, on the ventral sur-
face, much of the medial and lateral
margins of the nearly circular fora-
men praepalatinum. Anterolater-
al^ a short, broad process extends
ventrolaterally to meet the medial
surface of the palatal exposure of
the maxilla. Posterior to this pro-
cess it forms the anteromedial and
medial margins of the apertura
narium interna and continues
posteriorly as a narrow rod be-
tween the palatines to meet the
pterygoids. This posterior portion
extends slightly ventral to the
palatines which meet in the midline
dorsal to the vomer. The ridge thus
formed is highest anteriorly and
disappears posteriorly where the
vomer and palatines form an es-
sentially even surface. For the
anterior three-quarters of its
length, the vomer slowly narrows
posteriorly ; the posterior quarter is
again wider, though not as wide as
the anterior end.

The vomer possesses, on its dorsal surface, a pair of short, rather
stout processes; they are very near its anterior end in the Portland
skulls, but occupy most of the length of the greatly reduced vomer of the
Solenhofen form. Each process meets the vertical plate of the prefrontal
dorsally and the anteromedial edge of the palatine posterolateral^. The
latter contact is missing in the Solenhofen skull due to breakage of the
palatines; however, it was almost certainly present. Between the two
dorsal processes of the vomer there is a narrow groove, the sulcus vomeri.
The sulcus forms the ventral quarter of the fissura ethmoidalis.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

Os pterygoideum

The pterygoids are paired, more or less creseentic elements which
form most of the posterior half of the palatal surface of the skull. They
meet in a sagittal suture for much of their length and, on their ventral
surface, widely separate the palatines from the basisphenoid.

PORTLAND SPECIMENS

Anteromedially the pterygoid is
bounded by the palatine and lateral
to this it forms the posterolateral
margin of the large foramen pala-
tinnm posterius. Anterolateral^
there is a massive processus ptery-
goideus externus which has a thin
anterior portion along the posterior
palatine foramen and meets the
posterior end of the maxilla. The
major part of the process is later-
ally directed and its end is slightly
expanded dorsally and ventrally,
much as in Chelydra. Thus the
lateral margin of the pterygoid is
markedly concave anteriorly, and

SOLENHOFEN SPECIMEN

The anterior part of the ptery-
goid is greatly arched, so that the
deep median trough continues
posteriorly from the palatines.
There is almost no processus ptery-
goideus externus, although the
pterygoid does possess a short
transverse suture with the postero-
medial end of the maxilla. The
suture between the pterygoid and
palatine runs a short distance
posteriorly from the maxillary con-
tact and then becomes transverse,
extending to the midline. Due to
breakage of the palatine, the medial
portion of this suture is not present.

there is a fairly marked waist to The small oval foramen palatinum
the united pterygoids. posterius lies in the parasagittal

portion of the pterygoid-palatine
suture. A small nubbin, which
represents the remnant of a pro-
cessus pterygoidcus externus, ex-
tends ventrolateral^' from the
lateral margin of the pterygoid a
short distance posterior to the fora-
men palatinum posterius. The lat-
eral margin of the anterior half of
the pterygoid thus is essentially
straight and nearly parasagittal
with its anterior ends only slightly
more lateral than its center, and
there is no waist to the united
pterygoids.

60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Posteriorly the ventral surface of the pterygoid sends out a long
wing, somewhat curved, which lies along the ventral surface of the
processus articularis of the quadrate. This wing forms the posteromedial
margin of the large fenestra subtemporalis. Medial to the wings, the two
pterygoids are separated by the roughly triangular basisphenoid with
which they have posteromedial sutures. There is a slight ridge on the
ventral surface of the pterygoid which curves posteromedially and
posteriorly from the posterior end of the processus pterygoideus externus
to the posterior end of the pterygoid-basisphenoid suture ; it presumably
served for attachment of the pterygoid muscle.

Posteriorly, the pterygoid is in contact with the basioccipital. Just
anterior to their suture and lateral to the small ridge, the small foramen
posterior canalis carotid intcrni enters the pterygoid which completely
surrounds it. Between the carotid foramen and the lateral margin, the
pterygoid contains a large and deep posteroventral concavity. The floor
of this concavity forms the ventral margin of the large fenestra postotica.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

Along the medial portion of this The posterior margin of the

margin, the pterygoid has a short pterygoid is here concave, and does
suture with the anteroventral end not reach the anteroventral end of
of the exoccipital. the exoccipital. There is a small

foramen on the ventral surface of
each pterygoid slightly anterior to
the carotid foramen and medial to
the small ridge.

In lateral view the pterygoid appears rather small. Posterodorsally
it possesses a long suture with the quadrate, and more anteriorly a dorsal
one with the large epipterygoid. The crista pterygoidea is very low
anteriorly, but rises dorsally in the region of the foramen nervi trigemini
where it is notched to form the ventral margin of the foramen.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The crista meets the parietal at The crista is bounded postero-

either side of the foramen and is dorsally by the prootic. Both the
thus separated from the prootic by latter bone and the epipterygoid
the parietal and quadrate. reach the margin of the foramen

and thus separate completely the
pterygoid and the parietal.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

At the base of the crista pterygoidea, between the epipterygoid and
quadrate, there is an irregular depression as if those two bones possessed
an unossified connection lying lateral to the pterygoid.

The dorsal surface of the pterygoid is partly concealed in the
specimens, but much of its structure may be seen. Posteriorly it forms
much of the floor of the large cavum acustico-jugulare.

PORTLAND SPECIMENS SOLENHOPEN SPECIMEN

The prootic and the large proccs- There is no bony connection be-

sus interfenestralis of the opisthotic tween the opisthotic and the ptery-
are sutured to the posterodorsal goid. Presumably the prootic does
surface of the pterygoid. meet the dorsal surface of the

pterygoid, but the area cannot be

seen clearly.

Most of the medial portion of the pterygoid is overlain by the
basisphenoid ; the large rostrum oasisphenoidale reaches nearly, if not
actually, to the anterior margin of the pterygoid. The other bones which
articulate with the dorsal surface of the latter element are those described
in the lateral view. There is a wide sulcus cavernosus along the dorsal
surface of the pterygoid lateral to the basisphenoid and prootic and
medial to the epipterygoid, crista pterygoidea, and quadrate. The various
foramina are hard to see clearly, and the following description cannot
be considered certain in all respects. The canalis caroticus interims at
some point in its forward course leaves the pterygoid and enters the basi-
sphenoid.

PORTLAND SPECIMENS

It is connected, about midway
along its course, with the sulcus
cavernosus by a small foramen pro
ramo nervi vidiani which enters the
sulcus between the pterygoid and
the prootic.

Anteriorly the course of the vidi-
an nerve cannot be determined with

SOLENHOFEN SPECIMEN

The carotid canal is connected in
some fashion with the canal from
the foramen which may transmit
the vidian nerve. The latter opens
to the dorsal surface of the ptery-
goid well anteriorly, just lateral to
the rostrum oasisphenoidale.

There is no separate canalis nervi

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS

certainty. There appears to be a
small foramen in the pterygoid just
lateral to the rostrum basisphenoid-
ale which may transmit it ; the fora-
men could not be traced with a
bristle. In one specimen (R3164)
there is a small canalis nervi vidiani
running between the pterygoid and
epipterygoid from the region of the
foramen mentioned above to the
anterior end of the suture between
these bones ; there is no trace of it
in another specimen (R2914), and
the situation is not clear in the third
(E3163). A pair of small (nutri-
tive?) foramina occur on the
ventral surface of the pterygoids,

,near the lateral

margin

in the

region of greatest lateral constric-
tion.

SOLENHOFEN SPECIMEN

vidiani between the pterygoid and
the epipterygoid.

Os epipterygoideum

The epipterygoid is a fairly large plate rising dorsally from the
pterygoid to meet the processus inferior parietalis. Its ventral margin
is horizontal and smooth, but its dorsal suture is irregular. A thin postero-
ventral process extends towards, but does not quite reach, the quadrate.

PORTLAND SPECIMENS

The pterygoid and parietal send
narrow processes to meet each other
on the anterior margin of the fora-
men nervi trigemini and thus ex-
clude the epipterygoid from the
margin of this foramen. The
canalis nervi vidiani runs between
the anteroventral border of the
epipterygoid and the pterygoid in
at least one specimen (see the
description of the pterygoid con-
cerning the variation in this canal).

SOLENHOFEN SPECIMEN

The epipterygoid forms the an-
teroventral margin of the foramen
nervi trigemini, thus separating the
pterygoid from the parietal. Along
the margin of the foramen, there is
a pronounced lip which extends
laterally and bears a blunt ventral
process so that the slightly concave
lateral surface of the bone has a
marked groove anteroventral to the
lip.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

i i i
PTR EPT PPE

Fig. 5. Portland skull, British Museum R3164. Lateral view of skull fragment.
Abbreviations: EPT, epipterygoid. FNT, foramen nervi trigemini. ICA, incisura
columellae auris. PPE, processus pterygoideus externus. PTR, processus troehlearis.
V, vomer.

Elements of the Braincase and Otic Region

Os basisphenoidenm

The basisphenoid forms a small, roughly triangular element when
seen in palatal view. It is bounded anterolaterally by the pterygoids and
posteriorly by the basioccipital ; the anterior apex of the triangle is
rounded. A parasphenoid is not present as a distinct element, although
it is possible that the remnants of one are fused with the basisphenoid.

solenhofen specimen
The posterior end of the basis-
phenoid is essentially transverse. It
is not raised, but gives the im-
pression of a ridge due to the
marked difference in angle between
the ventral surfaces of the basis-
phenoid and the basioccipital.

PORTLAND SPECIMENS

The posterior end of the basis-
phenoid is very slightly concave,
and raised into a low transverse
ridge across the flat palatal surface
of the skull.

64 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Dorsally, the basisphenoid is more extensive and forms a considerable
part of the floor of the cranial cavity. Besides its lateral contact with the
pterygoids, which it also overlies anteriorly, and its posterior suture with
the basioccipital, the basisphenoid possesses an extensive dorsolateral
suture with the prootic. The dorsal surface can conveniently be divided
into two main portions separated by the dorsum sellae.

The more posterior portion forms a large trough which is slightly
concave medially. Posterolateral^ this surface forms part of the ventral
margin of the large hiatus acusticus or opening between the cavum cranii
and cavum labyrinthicum, thus separating the basioccipital from the
prootic. Each side of this dorsal surface is pierced by a small foramen,
the posterior opening of the ca7ialis nervi abducentis. The anterior open-
ing of this canal is into the medial side of the sulcus cavernosus.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

Anterior to the prootic suture, Anterior to the prootic suture,

the lateral margin of the basisphen- a prominent shelf on the basisphen-
oid continues anteriorly as the oid projects forward well dorsal to
processus clinoideus. This process the anterior portion of the bone,
extends a short distance dorsolater- The anterior margin of this shelf
ally as well as anteriorly, and ends shows a broad median extension
bluntly. Between the clinoid pro- flanked by two large concavities,
cesses of opposite sides, the basis- and, at the lateral margins, by two
phenoid slopes anteroventrally, the small projections or processus clin-
sloping region being the dorsum oidei. The dorsum sellae, between
sellae. the anterior and posterior parts of

the basisphenoid, is thus an essenti-
ally vertical face which is concave
anteriorly.

Laterally the basisphenoid enters the medial wall of the sulcus
cavernosus, thus separating the pterygoid and prootic anteriorly ; the sulci
converge anteriorly as the basisphenoid narrows. The anterior opening
of the canalis nervi abducentis lies in the wall of the sulcus well anteriorly
and just dorsal to the pterygoid-basisphenoid suture.

Anteriorly the basisphenoid forms a narrow projection, the rostrum
basisphenoidale, which lies on the dorsal surface of the pterygoids. This
rostrum terminates rather abruptly near the anterior end of the ptery-
goid ; quite possibly it was continued forward dorsal to the palatines as a
cartilaginous process. In transverse section the rostrum is formed by a
pair of oval bars of bone, one on either side with the long axis of the oval
horizontal. Between the ovals ventrally there is a very thin sheet of bone.
Thus there is a median groove in the dorsal surface of the rostrum ; the

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

posterior portion of this groove is the sella turcica. Although the entire
rostrum is more flattened in the Solenhofen skull than in the Portland
ones, the basic pattern is identical. At the posterior end of the sella
turcica, between the bases of the thicker lateral portions of the rostrum,
lie the entrances of the arterial circulation of the head, the foramen
anterior canalis carotid interni. The foramen is paired in the Portland
skulls, but appears to form a single median opening in the Solenhofen
skull. The canalis caroticus intcrnus must enter the basisphenoid from
the pterygoid ; the position and nature of this entrance are unknown.

Os basioccipitale

The basioccipital is a rather small but massive element at the postero-
ventral end of the skull. It makes up the greater part of the condylus
occipitalis, and forms all of the pit for the reception of a ligamentum
apicis dentis. The condyle lacks any pronounced neck and protrudes but
slightly.

PORTLAND SPECIMENS

The basioccipital is excluded
from the ventral margin of the
foramen magnum by the two ex-
occipitals which meet above it.

Ventrally the tubercula basioc-
cipitales are only slightly devel-
oped, and the crescentic precondyl-
ar fossa between them is shallow.
It does not extend anteriorly as far
as the basisphenoid.

SOLENHOFEN SPECIMEN

The basioccipital enters the floor
of the foramen magnum, narrowly
separating the two exoccipitals.

The tubercula basioccipitales are
large and prominent, as is the pre-
condylar fossa which extends an-
teriorly to the posterior margin of
the basisphenoid.

The basioccipital is bounded dorsally by the exoccipital and anterior-
ly by the pterygoids and the basisphenoid.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

Laterally the pterygoid and ex- Laterally the basioccipital enters

occipital are in contact thus ex- into the ventromedial margin of the

eluding the basioccipital from the fenestra postotica and cavum acu-

margin of the fenestra postotica stico-jugulare between its sutures

and the cavum acustico-jugidare. with the pterygoid and the exoc-

The exoccipital-basioccipital suture ciptal. The latter suture is irregu-

is straight. lar.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Dorsally, the basioccipital forms the posteromedian part of the
floor of the cavum cranii. There is a moderate-sized crista dorsalis basioc-
cipitalis at its anterior end in the Portland form ; this crista is much
reduced in the Solenhofen skull.

PORTLAND SPECIMENS

Due to the dorsal fusion of the
basioccipital with the exoccipitals,
the extent of the former cannot be
determined. In the largest skull
(R2914) both these elements ap-
pear to be excluded from the
margin of the hiatus acusticus ;
ihowever, in the small specimen
(R3164) both appear narrowly to
enter its postero ventral margin.

SOLENHOFEN SPECIMEN

The basioccipital makes only a
thin median band in the floor of
the cavum cranii; most of it is
overlain by the exoccipitals from
which it is separated by a very ir-
regular suture. Anteriorly, how-
ever, it is exposed laterally, and
narrowly enters the ventral margin
of the hiatus acusticus.

Os exoccipitale

Each exoccipital is a small element lying lateral to the foramen mag-
num. It is in two main parts, a dorsolateral and a ventromedial. The
dorsal portion is a rather narrow bar of bone lying posterior and ventro-
medial to the opisthotic. Dorsomedially, it possesses a short suture with the
supraoccipital and forms the dorsolateral margin of the foramen magnum,
and ventrolateral^ it lies along the dorsomedial margin of the fenestra
postotica.

From near the center of this dorsal bar, a strong process extends
ventromedially to the basioccipital. Medially this process forms the
ventrolateral margin of the foramen magnum, and laterally it enters into
the medial margin of the fenestra postotica.

PORTLAND SPECIMENS

The process expands laterally
where it meets the pterygoid
posterolaterally and possesses a
long suture with the ventral margin
of the opisthotic. Medially, the
process forms the dorsolateral
portion of the condylus occipitalis,
and appears to meet the opposite
exoccipital, thus excluding the basi-
occipital from the foramen mag-
num.

SOLENHOFEN SPECIMEN

The process is rather small, and
does not reach the pterygoid.
Medially, but not laterally, it pos-
sesses a very short suture with the
ventral margin of the opisthotic.
Medially, the process forms the
dorsolateral portion of the condylus
occipitalis. The basioccipital enters
the foramen magnum and thus
separates the two exoccipitals.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

At approximately the level of the top of the occipital condyle, the
ventromedial process of the exoccipital is pierced by two small foramina
nervi hypoglossi. One (larger in the Solenhofen skull; they are of
equal size in the Portland form) lies slightly posterodorsal to the other.
Anterolateral to these foramina, the exoccipital forms part of the margin
of the fenestra postotica.

PORTLAND SPECIMENS

The medial portion of this
fenestra is separated from the
lateral part by a ventral process
of the opisthotic and forms a canal
for the vena cerebralis posterior
and the vagus and accessory nerves.
Its posterior opening, the foramen
jugular x posterius, is bounded
dorsally, medially, and ventrally
by the exoccipital, with the opis-
thotic forming the lateral wall. At
its opening into the cavum cranii,
the foramen jugulare antcrius, it is
surrounded by the same two bones,
with the exoccipital forming its
dorsal, posterior, and ventral mar-
gins. The canal thus formed runs
somewhat dorsally as well as antero-
medially, and forms the anterior
border of the exoccipital. About
two-thirds of the way anteromedi-
ally along the canal there is a
prominent fenestra perilymphati-
ca; this lies mainly within the
opisthotic, but the exoccipital may
form part of its ventral margin.
The sutures in the area are not
perfectly clear and it is impossible
to tell whether or not such is the
ease.

SOLENHOFEN SPECIMEN

The medial portion of this fenes-
tra, that bordered by the exoccipi-
tal, forms a median notch for the
vena cerebralis posterior and the
vagus and accessory nerves (an in-
complete foramen jugulare posteri-
us). The canal for these structures
runs anterodorsomedially along the
anterior margin of the exoccipital
to enter the cavum cranii through
the foramen jugulare anterius.
This foramen is surrounded by the
exoccipital except anteriorly where
the opisthotic enters its margin.
Near the medial end of the canal,
the exoccipital forms the floor of
the large fenestra pcrilymphatica
which leads anteriorly through the
opisthotic. The latter forms its
medial, dorsal, and lateral margins.
Ventrolaterally the margin of the
foramen is incomplete, exactly like
the same margin of the foramen
jugulare posterius; in both cases,
the opisthotic and exoccipital just
fail to make contact ventrolaterally.

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOOY

FOV-

-ACS

PIF FJP

FNH

Fig. 6. Portland skull, British Museum E3164. Posterior view of skull fragment,
modified to show both cava acustico-jugulares at comparable posterolateral angles.
Abbreviations: ACS, aditus canalis stapedio-temporalis. CC, canalis cavernosus. FEG,
foramen externum nervi glossopharyngei. FJP, foramen jugulare posterius. FNH,
foramen nervi hypoglossi. FOV, fenestra ovalis. FPL, fenestra perilymphatica. PIF,
processus interfenestralis.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

ACS-n FEG

Fig. 7. Solenhofen skull, Teyler Museum. Posterior view. Abbreviations: ACS,
aditus canalis stapedio-temporalis. CC, canalis cavernosus. FEG, foramen externum
nervi glossopharyngei. FNH, foramen nervi hypoglossi. PIF, processus interfenestralis.

Os supraoccipitale

Much of the posterodorsal part of the supraoccipital is missing in all
the specimens. There was at least some vertical portion which rose to
meet the parietals ; however, whether the supraoccipital actually formed
any part of the dorsal roof or possessed any crista supraoccipitalis cannot
be determined.

The supraoccipital forms the roof of the posterior part of the cavum
cranii and of the otic region and the posteromedial portion of the floor
of the fossa temporalis where it is bounded anteriorly by the parietal,
anterolateral^ by the prootic, and posterolateral^ by the opisthotic. At
its posterior end the supraoccipital possesses a short suture with each
exoccipital and, between them, forms a narrow part of the dorsal margin
of the foramen magnum. Within the cavum cranii, the supraoccipital is
bounded by the same elements ; between its sutures with the prootic and
opisthotic it forms the dorsal margin of the large hiatus acusticus.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS

The details of the otic region can
be studied, in part, in the fragmen-
tary specimen (R3163). Apparent-
ly the supraoccipital forms very
little of the roof of the cavum
labyrinthicum ; the recessus laby-
rinthitis supraoccipitalis is a very
small cavity, probably containing
only the crus commune. It is repre-
sented by a depression in the
ventral surface of the supraoccipi-
tal. From this there is a canal
leading anteriorly into the prootic
for the canalis semicircularis an-
terior and another leading posteri-
orly to enter the opisthotic. The
latter contained the canalis semi-
circularis posterior. There appears
to be no foramen aquaducti vesti-
buli; a small notch is visible in the
dorsal rim of the hiatus acusticus,
but it mav be the result of wear.

SOLENHOFEN SPECIMEN

The details of the otic region are
obscure in the articulated skull.
Certainly there is within the supra-
occipital or neighboring bones a
well developed recessus labyrinthi-
tis supraoccipitalis. Nothing can
be seen of the semicircular canals.

Os opisthoticum

The opisthotic is a roughly triangular element, the dorsal side of
which forms the posterior end of the floor of the fossa temporalis.
Dorsally it is bounded anteromedially (along the shortest side of the
triangle) by the supraoccipital. The posterior edge possesses a suture
with the exoccipital medially, but laterally it enters the posterior margin
of the skull and the roof of the fenestra postotica, The exoccipital-opistho-
tic suture is long in the Portland skulls, but greatly shortened in the
Solenhofen specimen. Most of the anterolateral margin of the opisthotic
(the longest side of the triangle) is bounded by the quadrate; laterally
the opisthotic also is in contact with the squamosal, especially in the
Solenhofen skull. The anterior angle of the opisthotic just touches the
prootic, but does not extend anterior as far as the foramen stapedio-
temporale. There is a much smaller foramen at the spot where the
opisthotic, prootic, and quadrate meet; it could not be probed with a
bristle. The dorsal surface of the opisthotic slopes gradually from a more
dorsal position medially to a more ventral one laterally. Its posterior
margin is somewhat ventral to its anterior one in the Portland skulls,
but nut in the Solenhofen form.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

The lateral half of the ventral face of this element is a flat surface
roofing much of the cavum acustico-jugulare ; medially it becomes far
more complex. A prominent processus interfenestralis extends antero-
ventrally from the main plate of bone, separating the more anterior cavum
labyrinthicum from the more posterior canal for the vena cerebralis
posterior. The process is thus a flat plate lying in the transverse plane.
The small foramen externum nervi glossopharyngei pierces this process
well dorsally ; its posterior end is near the lateral margin of the process,
and its anterior near the center so that the opening runs laterally as well
as posteriorly. Near its lateral end the processus interfenestralis forms
the anterior margin of the foramen jugulare posterius and the posterior
margin of the fenestra ovalis, while medially the process forms the an-
terior boundary of the foramen jugulare anterius and the posterior
boundary of the hiatus acusticus.

PORTLAND SPECIMENS

The ventral end of the process is
sutured to the exoccipital medially
and to the pterygoid laterally.
Near its medial end, the ventral
margin is pierced by the large
fenestra perilymphatica ; whether
this fenestra is completely sur-
rounded by the opisthotic or
whether the exoccipital enters its
ventral margin cannot be deter-
mined. The anteromedial margin
of the processus interfenestralis is
notched ventrally, possibly for the
passage of the glossopharyngeal
nerve. The notch may thus be an
incomplete foramen internum nervi
glossopharyngei.

In E3163 the interior of the
cavum labyrinthicum may be seen.
There is, in the anterior face of
the opisthotic, a large posterodorsal
recess of the cavum labyrinthicum,
the recessus labyrinthitis opisthoti-
cus ; it is a roughly hemispherical
cavity. The canalis semicircular is
posterior leads from its postero-
dorsal wall into the supraoccipital.

SOLENHOPEN SPECIMEN

The ventral end of the process
meets the exoccipital. Lateral to
their contact there is a very large
notch for the fenestra perilym-
phatica. The ventrolateral end of
the process fails to reach the exoc-
cipital or pterygoid so that the
ventral margins of the foramen
jugulare posterius and fenestra
perilymphatica are incomplete.
There is a small foramen internum
nervi glossopharyngei in the antero-
medial margin of the processus
interfenestralis which leads from
the cavum labyrinthicum to the
cavum cranii.

Nothing can be seen of the inside
of the cavum labyrinthicum; as in
all turtles, the opisthotic here forms
its posterior wall.

72 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS

Anterolaterally there is a wider
foramen for the canalis semicircu-
laris horizontalis. The latter runs
anteriorly to enter the large reces-
sus labyrinthicus prooticus; the
canal is separated from the cavum
labyrinthicum only by a narrow bar
of the opisthotic and is not bounded
medially by the prootic at any
point.

Internally the opisthotic is bounded by the supraoccipital dorsallv
and the prootic anteriorly. Its ventral margin roofs the fenestra ovalis.

Os prooticum

The prootic is a fairly small, irregularly shaped element lying in
the anteromedial portion of the otic region. Its dorsal surface is a four-
sided plate forming the anteromedial quarter of the floor of the fossa
temporalis. Posteromedial^ it possesses a short suture with the supra-
occipital and medially an equally short one with the processus inferior
parietalis. Its longer anterior border forms the anterior margin of the
floor of the fossa temporalis, including at least half of the prominent,
spout-shaped processus trochlearis. There is a long posterolateral suture
with the quadrate. Near the posteromedial end of this suture, the quad-
rate and prootic are separated by the large foramen stapedio-temporale.
The foramen leads into a short but wide canalis stapcdio-temporalis which
runs ventrolateral^* between the same two bones to open into the dorso-
lateral corner of the cavum acustico-jugulare, immediately dorsal to the
posterior end of the canalis cavernosus.

PORTLAND SPECIMENS SOLENHOPEN SPECIMEN

The prootic forms roughly half The prootic forms nearly all of
to two-thirds of the moderately the exceedingly well developed pro-
large processus trochlearis. Antero- cessus trochlearis. Anteroventrally,
ventrally, it is bounded by the it is bounded by the quadrate and,
quadrate and has only a small ex- medially, by the pterygoid. The
posure on the surface of the skull, prootic enters the posterodorsal
The processus inferior parietalis margin of the foramen nervi trige-
sends a thin extension ventrally mini, and thus narrowly separates
which meets the pterygoid posterior the processus inferior parietalis
to the foramen nervi trigemini, thus from the pterygoid.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 73

PORTLAND SPECIMENS

excluding the prootic from the mar-
gin of that foramen. "Whether or
not the parietal and quadrate are
in contact and separate the ptery-
goid and prootic posterior to the
foramen cannot be determined with
certainty, bnt it seems probable.

The lateral surface of the prootic forms the dorsolateral wall of the
canalis caver nosus and of its anterior opening into the cavum cranii, the
foramen cavemosum. Near its ventral border it is pierced by the foramen
nervi facialis which enters the canalis cavernosus.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

Just ventral to this foramen in This foramen may be seen

R3163, the small foramen pro raino through the foramen magnum, but

nervi vidiani passes between the could not successfully be probed,

prootic and the pterygoid before Whether or not a foramen pro ramo

entering the latter bone to emerge nervi vidiani is present cannot be

in the canalis caroticus intern us. determined.

On its medial surface, the prootic possesses sutures with the basis-
phenoid ventrally, the supraoccipital posterodorsally, and the parietal
anterodorsally. Posteriorly it forms the anterior margin of the hiatus
acusticus and, anteriorly, its free surface is notched medial to the fora-
men nervi trigemini. Near its ventral end there is a conspicuous depres-
sion in the surface of the bone, the fossa acustico-facialis. The medial
end of the foramen nervi facialis lies in the antero-ventral corner of
this fossa.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

Posterodorsal to the foramen Near the posterior margin of the

nervi facialis, there is a single, fossa, there is a very small foramen,

fairly large foramen nervi aenstici. presumably a foramen nervi acusti-

Since the posterior margin of the ci. Other details of the medial

prootic is somewhat eroded, it can- surface of the prootic cannot be

not be definitely stated that there seen in this specimen. The posterior

was only one such foramen ; how- face of this bone is also mainly

ever, there is no evidence for any obscured. It possesses sutures with

others. In posterior view, the the basisphenoid ventromedially,

prootic possesses a ventromedial the pterygoid ventrolaterally, the

suture with the basisphenoid and a quadrate laterally, and the opistho-

ventrolateral one with the ptery- tic dorsolaterally. The last two

goid. Laterally it forms the dorso- sutures are separated by the aditus

74 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

medial wall of the canalis caverno- canalis stapedio-temporalis, and

sus and, dorsally, possesses a suture those with the pterygoid and the

with the quadrate. The last suture quadrate by the canalis cavernosus.

is interrupted by the aditus canalis The anterior margin of the fenestra

stapedio-temporalis. Dorsally, it ovalis is formed by the posterior

meets the opisthotic. edge of the prootic between its

Much of the posterior wall of the sutures with the basisphenoid and

prootic is hollowed out to form the opisthotic.

large recessus labyrinthicus prooti- The anterior wall of the cavum

cus. This recess receives antero- labyrinthicnm is formed by the

dorsally the canalis semicircular is prootic; it cannot be seen in this

horizontalis. The posteroventral specimen,
margin of the recess forms the
anterior and antero ventral borders
of the fenestra ovalis.

Os quadratum

This quadrate is a rather large, irregularly shaped element much
like that of modern turtles. Its triangular dorsal surface forms most of
the lateral half of the floor of the fossa temporalis. This surface is
bounded posteromedially by the opisthotic, anteromedially by the prootic,
and, along the posterior half of its lateral margin, by the squamosal. In
the Solenhofen form, the anterolateral margin of the quadrate is bounded
by another (unidentifiable) bone of the skull roof. However, in the
Portland form the lateral margin is not perfect on any of the specimens ;
whether or not it was joined to any of the dermal elements of the skull
roof cannot be determined. At its anteriormost extremity, the quadrate
forms about a third of (in the Portland skulls) or just barely enters (in
the Solenhofen) the lateral side of the processus trochlearis which is
described above.

The medial wall of the quadrate is concave, and forms the lateral
wall of the cavum acustico-jugulare posteriorly and of the large canalis
cavernosus anteriorly. Ventral to these cavities, there is a suture between
the pterygoid and the quadrate; dorsal to them, the latter meets the
opisthotic and prootic as on its dorsal surface. The canalis stapedio-
temporalis runs anteromedially as well as dorsally in the quadrate-
prootic suture from its aditus which opens into the cavum acustico-
jugulare to the foramen stapedio-temporale which opens into the fossa
temporalis. Anteriorly the quadrate is a fairly simple plate.

Along its dorsal margin, the quadrate forms the anterior edge of
the floor of the fossa temporalis laterally and possesses a suture with the

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

prootic medially. This suture passes along the underside of the processus
trochlearis so that the quadrate forms a considerable portion of the
ventral surface of the process. The anteromedial margin of the quadrate,
which is slightly anterior to its anterolateral margin, joins the pterygoid
in a moderately short vertical suture. This suture turns sharply
ventrolateral^ and continues along the medial half of the ventral margin
of the quadrate which thus rests on the posterior wing of the pterygoid.
Although the pterygoid separates the quadrate from the epipterygoid,
there is a groove along the surface of the pterygoid between these two
elements which may have contained a cartilaginous processus epiptery-
goideus of the quadrate. The quadrate does not enter the margin of the
foramen nervi trigemini.

PORTLAND SPECIMENS

Since the processus trochlearis is
not exceptionally large, the anteri-
or face of the quadrate is only
slightly concave, being nearly
vertical ventrally and sloping an-
teriorly near its dorsal end.

At its ventrolateral end, the
quadrate extends a short distance
ventrally as the processus articular-
is. The actual articulating surface
(condylus mandibularis) is pre-
served in only one specimen ; it is
a small parallelogram with the long
sides in the transverse plane and
the shorter ones running postero-
laterally at angles of approximately
twenty degrees from the sagittal
plane.

SOLENHOFEN SPECIMEN

With the extreme development of
the processus trochlearis, the an-
terior face of the quadrate is ap-
proximately vertical in its ventral
half, but turns sharply anteriorly
and is nearly horizontal dorsally.

At its ventrolateral end, the
quadrate forms a broad but very
short processus articularis. This
bears the large condylus mandibu-
laris, a rather long ovate area with
the long axis in the transverse
plane and the lateral end wider
than the medial. It is somewhat
arched, and its lateral and medial
ends are both ventral to its center.

In posterior view little of the quadrate can be seen. Part of its
exposure in the floor of the fossa temporalis is visible as is the processus
articularis. Between these two parts, the squamosal and opisthotic join
dorsal to the fenestra postotica and the incisura columellac auris; the
latter is described below.

In lateral view, most of the quadrate is occupied by the large cavum
tympani.

PORTLAND SPECIMENS

The cavum is a rather shallow
bean-shaped depression which is
deepest posterodorsally. At the

SOLENHOFEN SPECIMEN

The cavum is a deep, nearly
circular depression. Its greatest
depth is dorsally and posterodors-

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS

posterodorsal corner, there is a
small opening between the quad-
rate and the squamosal ; presum-
ably it led into an antrum postoti-

SOLENHOFEN SPECIMEN

ally where it extends posteriorly
into the squamosal as the antrum
postoticum. There is no distinct
boundary between the antrum and

cum, but most of the squamosal is the cavum tympani. The incisura

lost so that this cannot be deter-
mined. There is a deep and quite
wide incisura columellae auris
which forms the hilum of the bean-
shaped cavum. It expands almost

columellae auris is deep, extending
anterodorsally from the postero-
ventral margin of the quadrate to
the center of the cavum, but it is
very narrow. This slit-like incisura

to the center of the quadrate, as extends along the posteroventral
seen in this view, from the postero- surface of the quadrate from the
ventral margin. The incisura con- lateral margin to enter the lateral
tinues as a deep groove along the wall of the cavum acustico-jugu-
posteroventral margin to join the I are. Along the ventrolateral mar-
larger medial indentation of the gin of the incisura, there is a
quadrate which forms the lateral prominent ridge on the quadrate,
wall of the cavum acustico-jugu- so that its posteroventral surface is
lare. quite concave.

Columella auris

PORTLAND SPECIMENS

The columella is unknown.

SOLENHOFEN SPECIMEN

The columella is a rather
straight, thin rod of bone extend-
ing medially and somewhat ante-
riorly from the center of the cavum
tympani to the fenestra ovalis, be-
tween the prootic and opisthotic.
It lies in the incisura columellae
auris laterally and then extends
across the center of the cavum
acustico- jugular e. Medially its end
is expanded into an oval oasis col-
umellae which fills the fenestra
ovalis. The long axis of the oval is
posterodorsal to anteroventral. The
basis columellae is a very low coni-
cal structure with the shaft of the
columella arising from its center.
Since the columella does not extend
laterallv to the lateral surface of

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 77

SOLENHOFEN SPECIMEN

the quadrate, it was presumably
continued in cartilage as in mod-
ern forms.

Cavum acustico-jugulare

The cavum acustico-jugulare forms a rather large cavity in the
posteroventral surface of the skull. It is widely open posteroventrally
through the large fenestra postotica. This fenestra is bounded by the
exoccipital medially and dorsomedially, by the opisthotic dorsally, by the
quadrate laterally and ventrolaterally, and by the pterygoid ventrally.
In the Solenhofen form, but not in the Portland one, the basioccipital
enters its ventromedial margin between the exoccipital and the pterygoid.
The cavum itself is largely roofed by the opisthotic, with the quadrate
taking part laterally, the prootic anteriorly, and the exoccipital postero-
medially. Its floor is formed by the pterygoid medially and the quadrate
laterally ; the prootic enters the floor anteriorly. In the Solenhofen skull,
the pterygoid is somewhat emarginated posteriorly so that the floor is
less extensive than in the Portland specimens. The cavum, and also the
fenestra postotica, is essentially drop-shaped with the larger, rounded
end medial. Laterally it tapers gradually, and is continuous with the
comparatively narrow incisura columellae auris. This incisura extends
laterally as a groove within the quadrate to enter the posteroventral
margin of the cavum tympani.

The cavum acustico-jugulare is partially divided by the processus
interfenestralis of the opisthotic into two parts, a smaller posteromedial
and a larger anterolateral chamber.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

Ventrally the process, which ex- The process extends ventrally
tends anteroventrally from the and slightly anteriorly as a thin,
body of the opisthotic, is solidly nearly transverse plate of bone. It
fused to the floor of the cavum does not extend to the floor of the
along the length of the opisthotic- cavum, and thus the foramen jugu-
prootic suture. It is a thin, nearly lare posterius, fenestra perilympha-
transverse plate of bone. tica, and fenestra ovalis are incom-

plete ventrally. Presumably there
was a cartilaginous extension of the
process which completed the mar-
gins of these openings ; if this was
the case, then the pattern is almost
identical with that seen in the Port-
land skulls, and one description
will suffice for both.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

The smaller, posteromedial portion of the cavum acustico-jugulare
is a short and rather wide tubular cavity leading medially to open into
the cavum cranii. It lies between the exoccipital and the opisthotic. The
lateral end of this cavity is the foramen jugulare posterius and the medial
end, opening into the cranial cavity, is the foramen jugulare anterius.
Two smaller foramina enter the anterior (opisthotic) wall of the canal
from the cavum labyrinthicum : dorsolaterally the small foramen ex-
ternum nervi glossopharyngei and ventromedially the larger fenestra
perilymphatica. Thus this passage serves for the transmission of the
glossopharyngeal, vagus, and accessory nerves and a small vein (vena
cerebralis posterior = vena jugularis interna of mammals), and is homo-
logous with the jugular foramen of mammals.

FAC FNF CC

Fig. 8. Portland skull, British Museum E3163. Anterior view of skull fragment.
Abbreviations: CC, canalis cavernosus. CL, cavum labyrinthicum. CNA, canalis nervi
abducentis. CSH, canalis semicircularis horizontalis. CSP, canalis semicircularis
posterior. FAC, foramen anterior canalis carotiei interni. FAF, fossa acustico-facialis.
FNF, foramen nervi facialis.

The larger, anterolateral portion of the cavum acustico-jugulare has
three large openings which are separated from each other by the prootic.
Dorsomedially there is a large circular fenestra, ovalis which lies between

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 79

the prootic and the opisthotic. It opens into the cavum labyrinthicum
and is occupied by the basis columellae. From this, the shaft of the
columella extends laterally across the cavum acustico-jugulare and into
the incisure/, columellae auris. Lateral to the fenestra ovalis, between the
prootic and the quadrate, there is a smaller foramen, the aditus canalis
stapedio-temporalis. The arteria stapedialis passes dorsally through it to
reach the fossa temporalis. Finally, ventral to the two openings described
above, there is the large posterior end of the canalis cavernosas. This
canal runs posteriorly and somewhat laterally from the cavum cranii and
contains the vena capitis lateralis. It is bounded by the quadrate laterally,
the prootic dorsomedially, and the pterygoid ventromedially.

Cavum labyrinthicum

The cavum labyrinthicum forms a relatively large chamber contain-
ing the inner ear. It lacks, in the bony skull, a medial wall and is widely
open into the cavum cranii through the hiatus acusticus. Presumably in
life this hiatus was, as in recent turtles, almost entirely closed by carti-
lage. Although basically rectangular, the margin of the hiatus is very
irregular ; since this irregularity has been increased by erosion of the
skulls, the details of its form are not worth elaborate description. Dorsally
the hiatus is bordered by the supraoccipital, anteriorly by the prootic,
ventrally by the basisphenoid, and posterodorsally by the opisthotic.
Posteroventrally both the basioccipital and exoccipital also enter its
margin in the Solenhof en skull ; the same is true of one of the Portland
skulls (R3164), but in another larger one (R2914) the opisthotic appears
to extend ventrally to meet the basisphenoid and thus exclude both the
basioccipital and exoccipital from the margin of the hiatus.

The cavum labyrinthicum is bounded posteriorly and posterolateral^
by the processus interfenestralis of the opisthotic and anteriorly and
anterolaterally by the prootic. These same two bones form the postero-
lateral and anterolateral thirds of the roof of the cavum. Between them
and medial to them, the supraoccipital possesses a triangular exposure
forming the medial third of the roof. Finally the ventral surface of the
cavum is formed in large part by the pterygoid with the prootic and
basisphenoid entering its anterolateral and medial sides respectively ; their
exposure in the floor of the cavum is larger in the Solenhofen than in the
Portland form. In the latter, but not the former, the opisthotic also
enters this floor posterolaterally.

Besides the hiatus acusticus, there are five other openings into the
cavum labyrinthicum which can be seen in the specimens. Three of these,
the fenestra perilymphatica, foramen externum nervi glossopharyngei,
and fenestra ovalis, enter the cavum acustico-jugidare as described above.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

They lie, respectively, veutromedially on the posterior wall, dorsolaterally
on the posterior wall, and ventrally on the lateral wall of the cavum
labyrinthicum; the first two are surrounded (or nearly so) by the opis-
thotic while the last is between the opisthotic and prootic. The foramen
internum nervi glossopharyngci pierces the opisthotic between the cava
labyrinthicum and cranii, and lies near the center of the posterior margin
of the hiatus acusticus. Although present as a foramen in the Solenhofen
skull, it is represented only by a small incisure in the margin of the opis-
thotic in the Portland form ; presumably cartilage completed the margin
of this small foramen. The foramen nervi acustici also connects the cava
labyrinthicum and cranii, but it enters the anteromedial portion of the
former, piercing the prootic. It is apparently a single foramen in both
forms. Although there is no foramen aquaducti vestibuli, the supra-
occipital (dorsal) margin of the hiatus acusticus is slightly notched; pos-
sibly the notch represents this foramen, but it is impossible to determine
whether it is natural or a result of erosion of the skull.

PORTLAND SPECIMENS

The inside of the cavum laby-
rinthicum can be seen in only one
specimen (R3163) ; the following
description is based wholly on that
one. There are, extending out pos-
terolateral^, dorsally, and ante-
riorly from the dorsal part of the
cavum, three prominent recesses,
the recessi labyrinthici opisthoti-
cus, supraoccipitalis, and prooticus
respectively. They lie within the
bones whose names they bear. The
prootic recess, which presumably
contained the ampullae of the can-
ales semicircular 'es anterior and
horizontal is, is the largest; the
opisthotic recess which contained
the third ampulla is larger than the
supraoccipital recess. As in most
recent turtles, all three semicircu-
lar canals lie, at least partially,
within separate canals in the bones.
Thus the canalcs semicircular es an-
terior and posterior run from the

SOLENHOFEN SPECIMEN

Almost nothing can be seen of
the interior of the cavum labyrin-
thicum, and description is, there-
fore, not possible.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 81

PORTLAND SPECIMENS

recessus labyrinthicus supraoccipi-
talis to the reccssi labyrinthici
prooticus and opisthoticus respec-
tively, passing through the supra-
occipital and the prootic or opis-
thotic. The third canal, the canalis
semi circular is horizontalis, runs
from the opisthotic to the prootic
recess, but is surrounded only bv
the former bone.

Cavum cranii

The cavum cranii will be described but briefly; further details are
given in the descriptions of the individual bones that surround it. Only
that portion of the cavity posterior to the orbital region is considered here.
Thus the cavity extends from the foramen magnum at its posterior end to
the anterior margin of the processus inferior parietalis. Dorsally the
cavum is roofed by the supraoccipital posteriorly and the parietal an-
teriorly. This roof is somewhat arched, being lowest posteriorly, and it
is not pierced by any foramina.

The ventral surface of the cavum is considerably more complex.
Most of it is formed by the dorsal surface of the basisphenoid, but its
posterior quarter is composed of the basioccipital medially and the exoc-
eipitals laterally. The pterygoid enters the floor of the cavum anterolat-
eral^, along the ventral side of the sulcus cavernosas. For the posterior
three-quarters of its extent this floor is a simple trough. In the Portland
form, there is a moderately prominent crista dorsalis basioccipitalis at
the anterodorsal end of the basioccipital ; this crista is much reduced in
the Solenhofen skull. A small canalis nervi abducentis runs anteroven-
trally through the basisphenoid from near its center on either side to
enter the anterior part of the sulcus cavernosus. The posterior portion of
the cranial floor ends at the dorsum sellae.

Anterior to this, the floor, formed by the rostrum basisphenoidale,
continues as a horizontal trough, but at a lower (more ventral) level than
farther posteriorly. The dorsum sellae, where the change in level occurs,
is described in the section on the basisphenoid ; it is pierced by the fora-
men anterior canalis carotid interni where the arteria carotis interna
enters the cavum cranii. Lateral to the rostrum basisphenoidale there is
a deep groove, the sulcus cavernosus, in which the vena capitis lateralis
lies. Posterior to the canalis nervi abducentis, the vein passes through the
foramen cavcrnosum, which lies lateral to the anterior margin of the

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

prootic, and hence leaves the cavum cranii. Immediately anterior to the
dorsum sellae and foramen anterior canalis carotid interni is the sella
turcica in which rests the pituitary ; the sella is not markedly differenti-
ated from the remainder of the anterior half of the cranial floor.

Seven bones enter the lateral wall of the cavum cranii. Dorsally it is
formed by the ventral part of the supraoccipital and the processus in-
ferior parietalis. Its ventral half consists of the exoccipital, the opis-
thotic {processus interfenestralis), the prootic, and the epipterygoid, in

CNA-

pip

FNT

_J.

RBS-1 LFAC

Fig. 9. Portland skull, British Museum E3164. Anterior view of region of
dorsum sellae. Abbreviations: CNA, canalis nervi abducentis. FAC, foramen anterior
canalis carotici interni. FNT, foramen nervi trigemini. PIP, processus inferior
parietalis. RBS, rostrum basisphenoidale.

order from posterior to anterior. The crista pterygoidea enters the ex-
treme ventral margin of the anterior quarter of the lateral cranial wall.
This wall is penetrated by a series of foramina, mostly for the passage of
all but the four most anterior cranial nerves (the canal for the abducens
is more ventral and has already been mentioned). Posteroventrally the
hypoglossal nerve exits through two small foramina nervi hypoglossi
which are surrounded by the exoccipital. Slightly anterodorsal to these,

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 83

between the exoccipital and the opisthotic, lies the larger foramen jugu-
lar e anterius; through it pass the vagus and spinal accessory nerves as
well as a small vein to enter the cavum acustico-jugulare. The opisthotic
and prootic are separated by the large hiatus acusticus described above.
In the Portland form its posterior margin is notched and in the Solen-
hofen skull there is a small foramen in the opisthotic just posterior to it,
the foramen internum nervi glossopharyngei. The supraoccipital or dor-
sal margin of the hiatus may also have a small notch, presumably for the
endolymphatic duct ; this notch was probably made into a foramen aqua-
ducti vestibuli by the cartilage which, in life, closed the hiatus acusticus.
Both these notches and the hiatus enter the cavum labyrinthicum. Just
anterior to the ventral part of the hiatus, there is a marked depression
in the prootic, the fossa acustico -facialis. From this fossa a single foramen
nervi acustici leads posterolateral^ into the cavum labyrinthicum. Other
foramina for the acoustic nerve may have been present in the cartilage
mentioned above. Another foramen, the foramen nervi facialis, runs
anteroventrally as well as laterally from the fossa acustico -facialis, just
anteroventral to the foramen nervi acustici. It serves for the passage of
the facial nerve and enters the canalis cavernosas. Finally the large oval
foramen nervi trigemini lies well anteriorly, between the processus in-
ferior parietalis and the crista pterygoidea. In the Portland form the
foramen is completely surrounded by these two elements, but in the
Solenhofen skull they are separated anteriorly by the epipterygoid and
posteriorly by the prootic so that all four bones enter the margin of this
foramen. Through it pass the maxillary and mandibular branches of the
trigeminal nerve.

',--

Mandible

Os dentate

The dentary is a large element forming most of the anterior half of
the mandible. The long symphysis is solidly fused. Laterally the surface
of the bone is slightly roughened, and possesses a series of nutritive fora-
mina, most of them near its dorsal margin.

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The anterior end of the dentary The anterior end of the dentary

extends dorsally to form a gently is not upturned, and there are no

curved, but quite strong beak. On ridges on the triturating surface,

the lateral edge of the triturating However, the labial margin of that

surface, there is a prominent labial surface is quite sharp. The sym-

ridge which is quite wide and blunt physis is fully half the length of

posteriorly, but becomes narrower the entire dentary, thus forming a

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

PORTLAND SPECIMENS

and sharper anteriorly. Separated
from the labial ridge by a deep
trough, there is a well developed,
sharp lingual ridge. The lingual
ridges of either side meet near the
middle of the long symphysis, and
do not continue to the anterior end
of the jaw. There are numerous
small nutritive foramina within the
trough between the ridges. The

SOLENHOFEN SPECIMEN

large, relatively plane area which
does, however, possess a distinct
elongate median concavity occupy-
ing most of its length. There are
numerous, fairly large nutritive
foramina on the triturating sur-
face, mostly just inside the labial
margin. Medially, the splenial ex-
tends anteriorly nearly to the sym-
physis so that the sulcus cartila-
ginis meckelii is very short.

medial surface of the dentarj7, be-
tween the splenial and the sym-
physis, is deeply grooved by the
sulcus cartilaginis meckelii; the
sulci of opposite sides are continu-
ous around the posterior end of the
symphysis.

Posterolaterally, the jaws are broken, and the relationships cannot
be fully determined. In medial view, the posterior half of the dentary is
overlain dorsally by the coronoid and more ventrally by the splenial.
Ventral to the latter element, the dentary extends posteriorly for an un-
known distance ; this region is eroded and the sutures between the ele-
ments cannot be seen.

Os spleniale

The splenial is a relatively large, plate-like bone on the medial sur-
face of the mandible. It lies medial to the sulcus cartilaginis meckelii,
and, in the Portland form, also ventral to its anterior portion. Thus the
sulcus is really better termed a canalis cartilaginis meckelii for most of
its length.

PORTLAND SPECIMENS

The splenial is a trapezoidal
plate bounded dorsally bj^ the coro-
noid, posterodorsally by the prear-
ticular, ventrally by the dentary

SOLENHOFEN SPECIMEN

The splenial is a diamond-shaped
plate bounded anterodorsally by
the coronoid, posterodorsally by
the prearticular, posteroventrally

(and probably also by the angular bjr the angular, and anteroventrally

which is too eroded to be identi-
fied), and anterodorsally by the
sulcus cartilaginis meckelii. There
is a foramen, presumably entering
the Meckelian canal, near the an-
terior end of the splenial.

by the dentary. Dorsal to the an-
terior point of the diamond, which
nearly reaches the symphysis, is
the opening of the short sulcus car-
tilaginis meckelii.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS

COFh

-FM

Fig. 10. Solenhofen mandible, Teyler Museum. Dorsal view. Abbreviations:
COE, coronoid. FM, fossa meckelii. SP, splenial.

PRA

ANG

Fig. 11. Solenhofen mandible. Medial view of posterior end, left side. Abbrevia-
tions: ANG, angular. COB, coronoid. PEA, prearticular. SP, splenial. SUE,
surangular.

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Os coronoideum

SOLENHOFEN SPECIMEN

The coronoid forms most of the
very high and rather posteriorly
directed processus coronoideus,
thus having a considerable lateral
exposure. Medially it appears as
a rather long, narrow element
bounded anterodorsally by the den-
tary and posteroventrally by the
splenial and prearticular (with the
first anterior to the second). The
coronoid enters the triturating
surface as a narrow splint of bone
medial to the dentary; this exten-
sion reaches nearly to the symphy-
sis. Near the center of its medial
surface, the coronoid possesses a
small foramen, presumably enter-
ing the canalis cartilaginis meckelii.

The posteroventral end of the coronoid is deeply and quite widely
notched to form the anterodorsal margin of the fossa meckelii, and of the
canalis cartilaginis meckelii which continues anteriorly from this fossa.
Medial to the prearticular, a thin extension of the coronoid forms much
of the medial wall of this fossa.

PORTLAND SPECIMENS

The coronoid forms most of the
moderately prominent processus
coronoideus. Laterally it is exposed
only at the tip of that process, but
medially it appears as a large
roughly triangular plate. It is
bounded ventrally by the splenial
anteriorly and by the prearticular
posteriorly, and anterodorsally by
the dentary. Near the anterior
apex, there is a prominent foramen
leading ventrolaterally into the
bone, presumably to the canalis
cartilaginis m eckelii.

Os praearticulare

PORTLAND SPECIMENS

The prearticular is a plate of
bone lying on the medial surface of
the mandible ventral to the coro-
noid and posterodorsal to the splen-
ial. Ventrally it meets an uniden-
tifiable bone, probably the angular ;
this region is much eroded and
little can be definitely determined.
Posterodorsally it forms much of
the medial margin of the fossa
meckelii, although the coronoid
partly separates the prearticular
from the fossa.

SOLENHOFEN SPECIMEN

The prearticular is a rather long,
thin element extending posteroven-
trally from its suture with the ven-
tral margin of the coronoid. It is
bounded anteroventrally by the
splenial and, farther ventrally, by
the angular. Although in medial
view the prearticular appears to
form the medial wall of the fossa
meckelii, actually it is almost com-
pletely separated from that fossa
by the thin posteroventral exten-
sion of the coronoid which lies

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 87

SOLENHOFEN SPECIMEN

along its medial surface. Postero-
ventrally the end of the prearticu-
lar is deflected medially and slight-
ly expanded to form a surface
facing- posterodorsally and some-
what laterally. Although most of
this surface is covered by the
articular which rests on it and on
the surangular, its anteromedial
portion forms the medial end of the
area articularis mandibularis which
forms the mandibular articulation
with the quadrate.

Os articulare

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The articular is not preserved. The articular is a small triangu-

lar element bounded laterally by
the surangular and medially by the
prearticular ; its posterior border
is eroded. Its entire dosal surface
forms, with the prearticular, the
medial half of the area articularis
mandibularis which articulates with
the condylus mandibidaris of the
quadrate.

Os surangulare

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The surangular is not preserved. The surangular is a bone on the

lateral surface of the mandible
very similar to the prearticular in
its general relations. Anteriorly it
forms the posterior half of the lat-
eral wall of the fossa meckelii. Pos-
terior to the fossa, the surangular
becomes considerably thicker, pos-
sessing a medial extension which
forms the concave posterior wall of
the fossa. It also has a small lateral
process at this level, but most of

88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

SOLENHOFEN SPECIMEN

the lateral surface is eroded and its
boundaries cannot be determined.
The posterior end of the surangu-
lar expands to form nearly half of
the area articularis manclibularis,
the half which articulates with the
lateral portion of the extended con-
dylus mandibularis of the quadrate.
The articular bone lies medial to
the posterior end of the surangular.
There is a small gap between the
latter and the prearticular just an-
terior to the articular, and the
posteromedial wall of the fossa
meckelii is thus incomplete.

Os angulare

PORTLAND SPECIMENS SOLENHOFEN SPECIMEN

The angular is not preserved. The angular lies along the ventral

margin of the posterior half of the
jaw. This area is much eroded, es-
pecially laterally, and little can be
seen. Medially it lies posteroventral
to the splenial and anteroventral
to the prearticular. Its ventral
margin and lateral surface are not
preserved, but the latter was prob-
ably quite extensive.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 89

DISCUSSION: TURTLE SKULLS AND THE ORIGIN

OF TURTLES

The interest which attaches to a turtle skull of Upper Jurassic
age does not depend upon knowledge of the shell with which it
was once associated or upon accurate knowledge of its systematic
position and relationships. Interest in this case attaches to the
antiquity of the fossil itself. A Jurassic turtle skull will manifest
one of the grades of organization attained by the turtles during
that period. It may not be (is not at all likely to be) unspecial-
ized for that period, and one cannot be sure which are its special-
ized and which its primitive characters. Yet because of its relative
closeness in time to the oldest turtles, it could be hoped that it
would reveal a stage or step in the evolutionary line which
stretches from the most ancient turtles to those of the Recent.

We are therefore disappointed to discover that two skulls of
Upper Jurassic age tell us astonishingly little about the evolution
of turtles, next to nothing about features primitive for turtles,
and nothing at all that in any way points to the group from which
turtles have been derived.

Old as they are, these skulls are of a modernized turtle type.
There is no feature revealed by them that cannot be rather closely
matched in some group of turtles still living. Four characters
shared by the two Jurassic skulls are clearly primitive : the large
splenial, the presence of nasals (inferred but not definitely
shown in the Solenhofen skull), the widely exposed fenestra pos-
totica, and the large epipterygoids. These characters are not
found in combination in any later turtle, although each of them
may still be met with in one or another of the surviving families.
In most living turtles the splenial is absent or extremely reduced,
and the nasals quite absent. The Chelidae, however, have splen-
ials of moderate size (not indeed as large as in the Jurassic
forms) and most members of this same family have nasals also.
In most Recent turtles the fenestra postotica is partly or mostly
walled off by secondary plates of bone, and in many Recent forms
the columella is distally encircled by the quadrate, sometimes with
considerable thickness of bone behind it. In the Recent sea
turtles, on the other hand, the fenestra postotica is as widely ex-
posed as in the two Jurassic turtles and the distal end of the
columella is as loosely lodged in a notch of the quadrate as in
the Portland form, less enclosed than in the Solenhofen skull.
The epipterygoids are absent in both families of Recent Pleuro-
dira — the Chelidae and the Pelomedusidae — and small in many

90 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

cryptodires, but other cryptodirous turtles have the epiptery-
goids almost as large as in the two Jurassic forms.

Each of the two Jurassic turtles presents peculiarities that do
not seem to have any precise parallel in any other known turtle.

The palatal ridging of the Portland form is peculiar in that
the inner maxillary ridge is exactly at the choanal border and is
separated by a rather deep concavity from the tomial ridge. All
Recent turtles in which a ridge is present at the choanal margin
(e.g. the Batagur series of emydines and Podocnemis) have one
or more ridges between the choanal ridge and the tomial one.

It is similarly the palate of the Solenhofen form that is some-
what peculiar. This, in contrast to the Portland form, has a
strongly developed secondary palate. This is a specialized feature
repeatedly developed in turtles ; the Solenhofen palate is remark-
able only in the reduction of the vomer which, while not vestigial
or absent as in some Podocnemis, is completely concealed in
palatal view, the maxillaries and palatines having met in a long
suture beneath it. The closest approach to this condition is seen
in the pelomedusids, none of which, however, are similar in detail.

We have then in these Jurassic species the quite familiar
picture of more or less specialized primitive forms. They are not
and they should not have been expected to be idealized diagram-
matic ancestral types. They are end products in an early radia-
tion of the turtles, but a radiation that occurred after turtles had
assumed their definitive form — a form which they have kept
basically unchanged into the Recent, though they have played
infinite variations upon it.

The gap between these Jurassic turtles and the earliest known
turtles, those of the Upper Trias, becomes very much more im-
pressive now that these skulls are thoroughly known.

The shells of the Triassic turtles are very completely testudi-
nate. They differ somewhat from the shells of Jurassic to Recent
forms (which are actually somewhat simplified) but they are
primitive only in a temporal sense, not in the sense of leading
backward toward conditions and structures non-chelonian and
ancestral.

The skull of Proganochelys (the one form in which a skull is
known — Triassochelys is, we believe, a synonym) is quite differ-
ent in this respect. The magnificent material at the Museum fiir
Naturkunde at Stuttgart will be described in detail by Dr. K.
Staesche. However, notes taken by and photographs furnished to
one of us (E. E. Williams) while a Guggenheim Fellow in 1953
permit a few general remarks, in part confirming Jaekel's (1916)
observations, in part emending them.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 91

As Jaekel inferred, the narial opening was divided in Progano-
chelys, though his specimen did not show a complete septum.
This division was accomplished, however, not as he imagined by
a weakly calcified cartilaginous downward prolongation of the
nasal (Jaekel, 1916, p. 152) but by dorsal processes of the pre-
maxillae, the tips of which are inserted between the nasals much
as in Recent lizards or ancient procolophonids (Nycteroleter,
Procolophon, cf. Romer, 1956). In all other turtles, fossil and
Recent, except Kallokibotion 1 of the Cretaceous of Transylvania,
the nares are undivided.

The quadrate of Proganochelys is, as Jaekel pointed out, very
different from that of all other turtles for which this bone is
known. In place of the greatly developed cavum tympani and
incisura columellae auris, already so typically shown in the Jur-
assic turtles that we have described, there is only a slight incurv-
ing of the quadrate — a faint indication of things to come.
Apparently there was no notch whatsoever for the columella,
and the tympanum must have been attached just to the posterior
edge of the quadrate instead of being for the most part bounded
by that bone. The condition seems as primitive as in many coty-
losaurs and more primitive than in Diadectes.

The Stuttgart skulls show that the palate of Proganochelys
was in some respects more primitive than Jaekel believed. In
place of the "large tooth-bearing parasphenoid " of Jaekel 's
figure and text, the Stuttgart skulls show a distinct interptery-
goid vacuity. Jaekel's specimen showed crushing in this area,
and indeed he admits as much (1916, p. 175; see also his photo-
graph, Tafel 5B). The photograph of a quite uncrushed skull
(PL 5) shows the actual situation clearly; a cultriform process
is plainly visible between the separated pterygoids. Posteriorly
also the situation is primitive ; the quadrate ramus of the ptery-
goid does not send any flange inward to floor the cranioquadrate
passage as in all other known turtles, and the foramina for the
vena capitis lateralis, the internal carotid, and the stapedial
artery, as well as the fenestra ovalis are all exposed in ventral
view. In all respects Proganochelys approaches the primitive
reptilian condition and is separated by a very sizable morpho-
logical gap from such modernized turtles as the Jurassic ones
we have described.

lExcept in this regard, this peculiar, relatively Late germs is not primitive. The
material (seen in the British Museum) is poorly preserved, the sutures difficult
to trace, and not made easier to see by the white paint with which Nopcsa marked
them (sometimes changing his mind and incompletely removing a previously
painted line). In certain places breaks and injury to the surface account for the
peculiar course of some of the cranial sutures shown by Nopcsa (1923).

92 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Currently the Jurassic families and many later turtles includ-
ing some that survived to the Pleistocene (Meiolania) are united
with the forms of the Trias in a suborder Amphichelydia. It is
quite clear from what has been said above that the morphological
gap is much greater between Triassic forms and all later ones
than within the latter group. If major divisions below the order
are to be made, the basic one obviously must be between the
Triassic ones and all the others.

Indeed, Proganochelys fits Watson's (1914) concept of "Ar-
chichelone" better than Eunotosaurus, which Watson presented
as a possible testudinate ancestor.

Seeley in 1892 and Watson much more elaborately in 1914
suggested Eunotosaurus africanus as a turtle ancestor. Watson's
procedure was especially interesting in that he reasoned from the
apparent agreement in many features of Eunotosaurus and a
hypothetical "Archichelone" or theoretical turtle ancestor con-
structed by extrapolation from the characters of the turtles then
known. His summary of the resemblances between Eunotosaurus
and "Archichelone" was more plausible in 1914 than it appears
to be now. The advance of our knowledge, both of living and
fossil turtles, and especially the discovery and description by
Jaekel (1916) of Triassochelys only shortly after the appearance
of Watson's paper, has, not surprisingly, required some revision
of his ideas. For the most part these revisions indicate greater
primitiveness for "Archichelone" than Watson was quite pre-
pared to admit.

We may with more confidence comment on these points since
one of us (E. E. Williams) in 1952 and 1953 while visiting
Europe as a Guggenheim Fellow examined the material of Euno-
tosaurus at the British Museum only a few months before seeing
the fabulously fine Triassic turtles at Stuttgart. The latter were
only briefly studied, but we may in consequence use the descrip-
tion by Jaekel of Triassochelys with greater confidence.

It will be useful to compare Watson's concept of "Archiche-
lone" with a more modern version of the same concept based on
Proganochelys-Triassochelys and with the verifiable characters
of Eunotosaurus. An examination of the British Museum Eunoto-
saurus leads one to sympathize with the observation of Broom
(1941), who stated: "I have examined the specimens in the
British Museum and three or four other specimens which are a
little better preserved than these, but I have never published any
descriptions of them, always hoping that a really good specimen
will be found."

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 93

We will follow the procedure of first stating Watson's predic-
tion of the characters of "Archichelone," following this with a
version modified in accordance with more recent knowledge, and
then compare the latter with Eunotosaurus.

skull. Watson: " 'Archichelone' had a roofed skull with a
primitive Reptilian palate, teeth, probably only a squamosal of
the temporal series and no facial (internarial) processes of the
premaxillaries ..."

Revised statement. Watson's summary holds good with minor
modifications. Jaekel (1916) has, somewhat doubtfully, described
a supratemporal in Triassochelys. Olson (1947 and 1950) would
prefer to call the questioned bone in Triassochelys an intertem-
poral. In all later turtle types only one bone of the temporal
series, usually regarded as the squamosal, is ever present. The
absence of internarial processes also does not antedate the defini-
tive differentiation of turtles. They were present in the oldest
turtles, as reported by Jaekel. Two skulls of Proganochelys also
show this internarial septum (originals seen at Stuttgart, a cast
in the British Museum). Nopcsa (1923) has described a complete
internarial septum in the much later (Cretaceous) turtle Kallo-
kibotion. One of us (E.E.W.) has seen the skull of this form in
the British Museum and can confirm Nopcsa 's observations.

Eunotosaurus. Extremely little of the skull is known. Portions
of the jaws and of the palate x are preserved, in each case very
imperfectly. Except for the small size of the teeth, described by
Watson as very small on the pterygoid, and only slightly larger
on the maxilla and premaxilla, the palate so far as can be seen is
merely primitive without special resemblance to that of turtles.
Even in the case of the teeth there is a significant contrast. As
appears from Jaekel's (1916) description, the teeth were ex-
tremely small on the maxilla and premaxilla in Triassic turtles
and somewhat larger and more numerous on the pterygoid and
vomer. As Broom (1941) has stated there are certainly more
teeth on the maxillae of Eunotosaurus than Watson described.
Watson believed that there was no internarial septum in Euno-
tosaurus.

neck. Watson: " 'Archichelone' must have had a long and
flexible neck. ... As all known chelonians have eight cervical
vertebrae, we are justified in claiming this number for 'Archi-
chelone'."

iSo little of the palate is visible that we would not ourselves choose to say
auything about it.

94 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Revised statement. The neck of 'Archichelone' was short, ar-
mored, and had no more than seven vertebrae. Jaekel (1916) has
described a well-preserved cervical column in Triassochelys. In
this the seven cervicals were short and amphicoelous with high
spines, two-headed ribs, and large dermal ossicles in the skin
above them. The eighth vertebra was still part of the trunk re-
gion and was added to the neck only in later types. This neck
could have had only a very limited flexibility. Truly flexible and
elongate necks did not evolve in turtles before the Cretaceous.

Eunotosaurus. In the one specimen of this form in which the
skull is preserved, it is found with its ventral surface applied to
the ventral surface of the trunk region, its anterior end pointing
backward. Watson believed that this position was only possible
if the neck were fairly long and flexible. To the extent that this
conclusion is justified, we must regard Eunotosaurus as more
specialized in the Permian than any turtle was before the Creta-
ceous. In fact, however, we know nothing about the neck of
Eunotosaurus.

trunk. Watson: " 'Archichelone' had a series of dorsal scutes
comparable to those of a crocodile. ... It is by no means unlikely
that it resembled many members of that group in having ventral
scutes not homologous with abdominal ribs. ' '

Watson further believed that "Eight of ten dorsal vertebrae
and their ribs have special relationships with dorsal and dorso-
lateral scutes, while the peripherals probably correspond to
another set of bony scutes." Nuchal and pygal were probably
originally connected with additional vertebrae in front of and be-
hind the specialized ones and "lost this primitive connection
after the formation of the shell."

Revised statement. Again certain corrections or qualifications
need to be made. Many authorities believe that the major portion
of the turtle plastron represents fused abdominal ribs. Jaekel,
indeed, believed that signs of this origin were still present in
Triassochelys ; at all events in this genus a subdivision of the plas-
tron into the relatively large paired units of later forms was not
visible. Again while eight is the usual number of vertebrae in-
volved in the shell in modern turtles, nine pleurals occur in a
number of fossil forms including the oldest well-known types,
Triassochelys = Proganochelys. One or more suprapygals occur
in addition to these nine. On the other hand, the eighth cervical
vertebra was fused to the nuchal in Proganochelys just as Watson
anticipated might be the case in "Archichelone."

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 95

Eunotosaurus. There were ten dorsal vertebrae. Watson de-
scribed them as follows : "these are all fairly similar in structure.
The first is short, the second somewhat longer, and the third very
long. The fourth, fifth and sixth are about as long as the third,
and the seventh to tenth show progressive diminution in length.
The structure of the individual vertebrae is best shown in the
type specimen. The centrum is very slender, particularly in the
fourth to seventh dorsals, and somewhat hour-glass shaped ; it is
completely pierced by the notochordal canal. The rib-facet is
carried on a very low and small process which in the mid-dorsal
region is placed at the extreme anterior end of the centrum,
whilst anteriorly and posteriorly it travels back to the middle of
its length. The neural arch is rather massive when compared
with the centrum but still very narrow. It bears very narrow
zygapophyses which seem to interlock strongly. The upper sur-
face of the neural arch is essentially flat, being represented only
by a low median ridge. The whole arch appears to be placed
very far forward on the centrum and may overlap its anterior
end. There is no definite transverse process but the ribs seem to
touch the sides of the neural arch.

' ' The first dorsal rib is a narrow, slightly bent and quite short
bone of an ordinary character. The remaining dorsal ribs, from
the second to the ninth, are of an extraordinary character. Each
is strongly curved, articulates by a facet on its proximal end with
the facet on the centrum, and then rises, until its upper surface
comes in contact with the neural arch ; there is no definite tuber-
culum and the capitulum is extremely feeble. The rib then rises
above the level of the neural spine so that the dorsal surface of
the vertebral column lies at the bottom of a groove formed by the
proximal ends of the ribs. The rib now turns outwards and down-
wards. The ribs widen very rapidly from the capitulum, so that
until just at the point where they turn downwards their lateral
borders actually touch. They are of a massive character through-
out, and each is strengthened by the development of a ridge along
its visceral surface. ' '

There is in addition to these structures perhaps a dorsal bony
armor above them, as Watson inferred from one specimen. Wat-
son says : " In this individual in the anterior dorsal region a small
patch of what is undoubtedly bone substance is shown lying a
considerable distance above the ribs and neural arches. This is
clearly divided into pieces, one of which forms a small round
ridge in the middle line. The rest of the patch shows a dividing

96 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

line running longitudinally and a transverse division apparently
coincident with the line where the two ribs below it meet. At the
posterior end of the specimen, at the same distance above the ribs,
a narrow line of bone is seen in transverse section. This specimen
gives conclusive evidence of the actual presence of dermal ossifi-
cations, but these are so incompletely preserved as to make any
statement of their distribution of very slight value. There seem,
however, to have been a median series and lateral rows. ' '

Broom (1941) describes the ribs a little differently: "The
upper half of each rib is broad and convex above, but with a deep
median keel, so that in section each rib is triangular. Further in
most specimens the edges of the ribs do not meet but each rib
overlaps the rib behind it. Then each rib has a long pointed lower
half which passes downwards and slightly inwards. Each rib
is singleheaded. "

That there is here a parallel to the turtle condition is undoubted-
ly true ; that it is a close parallel we do not believe. The semblance
of a carapace is achieved by a broadening of the ribs them-
selves, so that they are already very wide close to their vertebral
articulations. In all true turtles including Proganochelys the ribs
themselves are not widened at all. From their articulations with
the vertebrae to their point of entrance into the carapace they are
not visibly very different from the ribs of more typical verte-
brates. Their apparent width in the carapace and their close
union with one another result from the addition to them of a
dermal component in which they are embedded. The lack of
width of the ribs themselves appears especially clearly when, as
in several lines of. marine turtles, the dermal component of the
carapace is strongly reduced and the ribs appear as separate,
narrow, well-spaced elements.

The pleural bone of a turtle is thus not pure cartilage bone and
not merely a rib ; it is the endochondral rib plus membrane bone
ossified around it. Ruckes (1929) in a study of the development
of the turtle carapace showed that the carapace is early repre-
sented by a thickening of the dermis with which the ribs and
neural arches secondarily gain connection and which they organ-
ize into pleurals and neurals. The ribs and neural arches give a
segmental character to the carapace, and they add a very firm
internal support, but the carapace is, according to Ruckes, inde-
pendent in its origin from the ribs and neural arches — originally
a quite separate dermal thickening. Ruckes' observations have
been confirmed by Walker (1947).

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 97

It is true that the situation in turtles is not simple. As Zan-
gerl's (1939) study has emphasized, there are two layers of
dermal ossification possible in turtles ■ — a deep thecal layer and
a superficial epithecal layer, but it is hardly possible to evade
the conclusion that both layers are dermal1 and that even the
deeper one is in no sense part of the ribs. In addition, the epi-
thecal layer seems to be a late development phyletically, unknown
before the late Cretaceous.

In regard to the elongation of the dorsal vertebrae in Eunoto-
saurus, thought by Watson to be a resemblance to "Archiche-
lone," it must be commented that the elongation is extreme in
Eunotosaurus, greater than in many modern turtles and much
greater in the middle Permian animal than in Proganochelys of
the late Triassic.

limb girdles. Watson : " In ' Archichelone ' the pectoral girdle
was undoubtedly in the usual position overlying the first dorsal
ribs, and must have been narrow relative to the bulk of the trunk
to allow of its passage back within the ribs. . . . The pelvic girdle
must also have been very narrow."

Revised statement. Undoubtedly in the remoter ancestors of
turtles the relation of shoulder girdle and ribs must have been of
the normal vertebrate type, but the hypothesis that the girdles
must have been narrow reflects a prior hypothesis as to the method
by which the change in that relation came about. It may or may
not be true.

Eunotosaurus. The limb girdles are primitive and small, but
not especially prophetic of conditions in turtles.

limbs. Watson: "We may take it as probable that 'Archiche-
lone' had limbs more or less like Eryops or Varanasaurus or
Dimetrodon." He mentioned, however, the narrow distal end of
the humerus and the short powerful downwardly directed radial
and ulnar crests of turtles and regards the presence of a shell as
reasons for these. The rounded and upturned head of the hu-
merus he described as "an obvious adaptation to the retraction
of the limbs."

Revised statement. Von Huene (1926) has described a Triassic
turtle humerus. It is much more primitive than any modern
type, especially in the greater width of the distal end and the

lit has been suggested that the deeper layer is periosteal (Goette, 1S99), but
in fact the thickening of the dermis which is to include both layers of carapace
bone occurs very early and is not a development of the perichondrium or peri-
osteum.

98 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

general shortness and stoutness of the bone. It is otherwise, how-
ever, very evidently a turtle humerus with a head and ulnar and
radial crests of a quite typical sort.

Eunotosaurus. The limbs are incompletely preserved but "the
development of a short powerful ulnar crest on the humerus and
the slight upturning of its head" are felt to be resemblances to
turtles. It should be noted, however, that, as Watson feels these
characters go along with the presence of a shell, they are resem-
blances implying affinity to turtles only if the shell truly indi-
cates such an affinity.

To sum up, Eunotosaurus does show a number of resemblances
to the turtles, but these resemblances are often to advanced types
instead of to early and primitive members of the order. The
apparent prototype of a chelonian carapace found in Eunoto-
saurus is in reality basically different in plan and is primarily
interesting as another method by which a similar result may be
achieved. Finally, the relation of limb girdles to ribs in Eunoto-
saurus is merely primitive and helps no more than do similar con-
ditions in other groups in visualizing the way in which the strange
condition so fundamental for turtles may have come about.

In thus dismissing Eunotosaurus we do not intend to disparage
Watson's shrewdness and insight in estimating the characters of
a turtle ancestor. We desire only to point out that in all its
characters, except its shell, Proganochelys fits his concept or is
more primitive than he felt justified in suggesting. In skull, neck,
even limbs and girdles we are not justified in asking for a more
primitive proto-turtle than the Triassic animals already provide.
We may indeed associate the proto-turtle with that broad and
varied group called Diadectomorpha, but it is at least the au-
thors' feeling that an attempt to derive the turtles directly from
any of the other diadectomorph groups fails, that the lineage of
turtles goes down alongside the other lineages to somewhere close
to the roots of the larger group itself. Turtles are, in fact, co-
ordinate with the other diadectomorphs rather than derivative
from them.

The shell of turtles and in particular its relation to the shoul-
der girdle remains an unsolved problem. It was this problem
which Watson attempted to settle by utilizing Eunotosaurus as a
turtle ancestor ; in this, we believe, he did not succeed.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 99

TERMINOLOGY

Unfortunately there is, at present, no standard system of ana-
tomical terminology in general use for turtle skulls ; almost
every author who has described them has invented his own names
for many structures. In the present work we have attempted to
devise a fairly consistent terminology, using familiar terms when-
ever they are appropriate and coining new ones only when none
of the previously proposed names seemed satisfactory. The fol-
lowing lists present the terms we have used and some of the more
familiar of their synonyms. Four earlier papers (Siebenrock,
1897; Kesteven, 1910; Ogushi, 1911; and Nick, 1912) are out-
standing in the detailed description of turtle skulls, and all their
terms equivalent to those used in the present paper are included.
It will be noted that even in the case of the most familiar struc-
tures there has been little uniformity in the names used by these
four authors. Other works are cited when they proposed or
standardized the use of a term or a system of terms which is not
used in any of the four major papers but which we have adopted.
In many cases the authors cited were not the originators of the
terms with which we have credited them. All those names for
which no author is cited are original. This rather elaborate list-
ing of terms is made in the hope, first, that it will be useful to
future workers comparing the discussion in the previous papers
on turtle skulls and, second, that it may bring about some stand-
ardization of such terminology.

T.erms Used in the Present Work:

aditus canalis stapedio-temporalis

antrum postoticum (Williams, 1954)

apertura narium externa (Siebenrock, 1897; Ogushi, 1911)

apertura narium interna (Siebenrock, 1897)

area articularis mandibulars

basis columellas (Siebenrock, 1897)

canalis alveolaris superior (Ogushi, 1911)

canalis caroticus internus (Siebenrock, 1897)

canalis cartilaginis meckelii

canalis cavernosus (Siebenrock, 1897; Nick, 1912)

canalis nervi abducentis (Siebenrock, 1897)

canalis nervi vidiani (Siebenrock, 1897)

canalis semicircularis anterior (Kesteven, 1910; Ogushi, 1911; Nick, 1912)

canalis semicircularis horizontals (Siebenrock, 1897)

canalis semicircularis posterior (Kesteven, 1910; Ogushi, 1911; Nick, 1912)

canalis stapedio-temporalis (McDowell, pers. comm.)

100 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

cavum acustico-jugulare (Kesteven, 1910)

cavum cranii (Nick, 1912)

cavum labyrinthicum (Ogushi, 1911)

cavum tympani (Siebenrock, 1897)

columella auris (Kesteven, 1910; Ogushi, 1911; Nick, 1912)

condylus mandibularis (Siebenrock, 1897)

condylus occipitalis (Siebenrock, 1897; Nick, 1912)

crista dorsalis basioccipitalis

crista pterygoidea (Siebenrock, 1897; Ogushi, 1911)

crista supraoccipitalis (Siebenrock, 1897; Nick, 1912)

dorsum sellae (Ogushi, 1911; Nick, 1912)

fenestra ovalis (Nick, 1912)

fenestra perilymphatica (Ogushi, 1911)

fenestra postotica (Nick, 1912)

fenestra subtemporalis (Bomer, 1956)

fissura ethmoidalis (Kesteven, 1910)

foramen alveolare superius (Siebenrock, 1897)

foramen anterior canalis carotici intend

foramen aquaducti vestibuli (Siebenrock, 1897)

foramen cavernosum (Nick, 1912)

foramen externum nervi glossopharyngeal (Nick, 1912)

foramen internum nervi glossopharyngei (Nick, 1912)

foramen jugulare anterius (Siebenrock, 1897; Nick, 1912)

foramen jugulare posterius (Siebenrock, 1897)

foramen magnum (Kesteven, 1910; Nick, 1912)

foramen nervi acustici (Siebenrock, 1897)

foramen nervi facialis (Siebenrock, 1897)

foramen nervi hypoglossi (Siebenrock, 1897)

foramen nervi trigemini

foramen orbito-nasale (Ogushi, 1911)

foramen palatinum posterius (Siebenrock, 1897)

foramen posterior canalis carotici interni

foramen praepalatinum (Seydel, 1896)

foramen pro ramo nervi vidiani (Siebenrock, 1897)

foramen stapedio-temporale

fossa acustico-facialis (Nick, 1912)

fossa meckelii (Siebenrock, 1897)

fossa nasalis

fossa temporalis (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

hiatus acusticus (Kesteven, 1910)

incisura columellae auris (Nick, 1912)

meatus choanae (Kesteven, 1910)

os angulare (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

os articulare (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

os basioccipitale (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

os basisphenoideum (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os coronoidum (Siebenrock, 1897; Nick, 1912)

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 101

os dentale (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

os epipterygoideum (Siebenrock, 1897; Kesteven, 1910; Nick, 1912)

os exoccipitale (Ogushi, 1911; Nick, 1912)

os frontale (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os jugale (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

osmaxillare (Siebenrock; 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os nasale (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os opisthoticum (Kesteven, 1910; Ogushi, 1911; Nick, 1912)

ospalatinum (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os parietale (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os postorbitale (Romer, 1956)

os praearticulare (Romer, 1956)

os praef rontale (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os praemaxillare (Siebenrock, 1897; Kesteven, 1910)

os prooticum (Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os pterygoideum (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os quadrato-jugale (Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os quadratum (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os spleniale (Romer, 1956)

os squamosum (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

os supraoccipitale (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

os surangulare (Romer, 1956)

processus articularis (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

processus clinoideus (Siebenrock, 1897; Kesteven, 1910; Nick, 1912)

processus coronoideus (Ogushi, 1911)

processus epipterygoideus (Siebenrock, 1897)

processus inferior parietalis (Ogushi, 1911)

processus interfenestralis (Kesteven, 1910)

processus pterygoideus externus

processus trochlearis (Schumacher, 1954)

recessus labyrinthicus opisthoticus

recessus labyrinthicus prooticus

recessus labyrinthicus supraoccipitalis

rostrum basisphenoidale (Zangerl, 1953)

sella turcica (Ogushi, 1911)

sulcus cartilaginis meckelii (Siebenrock, 1897; Nick, 1912)

sulcus cavernosus (Siebenrock, 1897; Ogushi, 1911; Nick, 1912)

sulcus olfactorius (Kesteven, 1910)

sulcus vomeri (Ogushi, 1911)

tuberculum basioccipitale (Siebenrock, 1897; Nick, 1912)

vomer (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912)

Terms Used in Other Major Works:

ampulla canalis seniicircularis frontalis (Siebenrock, 1897) = recessus labyrinthicus

opisthoticus
ampulla canalis seniicircularis sagittalis (Siebenrock, 1897) = recessus labyrinthicus

prooticus

102 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

antivestibulum (Ogushi, 1911) = cavum acustico-jugulare (In part)
antrum mastoideum (Siebenrock, 1897) = antrum postoticum
canaliculus pro nervo abdueente (Ogushi, 1911) = eanalis nervi abducentis
canaliculus pro ramo communicante n. palatini (Ogushi, 1911) = foramen pro ramo

nervi vidian!
eanalis abducens (Kesteven, 1910) = eanalis nervi abducentis

eanalis arteriae temporalis posterior (Ogushi, 1911) =: eanalis stapedio-temporalis
eanalis caroticus (Kesteven, 1910; Ogushi, 1911; Nick, 1912) = eanalis caroticus

internus
eanalis caroticus externus (Siebenrock, 1897) = eanalis stapedio-temporalis
eanalis deuterencephalicus (Ogushi, 1911) — cavum cranii (in part)
eanalis facialis (Kesteven, 1910; Ogushi, 1911; Nick, 1912) = foramen nervi facialis
eanalis glossopharyngeus (Kesteven, 1910) = foramina externum et internum nervi

glossopharyngei
eanalis hypoglossi (Kesteven, 1910; Nick, 1912) = foramen nervi hypoglossi
eanalis inframaxillaris (Ogushi, 1911) = eanalis cartilaginis meckelii
eanalis jugularis (Kesteven, 1910) = eanalis cavernosus
eanalis olfactorius (Ogushi, 1911) = sulcus olfactorius

eanalis palatinum major et minor (Ogushi, 1911) = foramen palatinum posterius
eanalis semicircularis externus (Nick, 1912) = eanalis semicircularis horizontalis
eanalis semicircularis frontalis (Siebenrock, 1897) = eanalis semicircularis posterior
eanalis semicircularis inferior (Kesteven, 1910) = eanalis semicircularis horizontalis
eanalis semicircularis lateralis (Ogushi, 1911) = eanalis semicircularis horizontalis
eanalis semicircularis sagittalis (Siebenrock, 1897) = eanalis semicircularis anterior
cavitas glenoidalis (Ogushi, 1911) = area articularis mandibularis
cavum archencephalicum (Ogushi, 1911) = cavum cranii (in part)
cavum intermedium (Ogushi, 1911) = cavum acustico-jugulare (in part)
cavum nasi (Kesteven, 1910) = fossa nasalis
cavum tympani (Nick, 1912) = cavum tympani, antrum postoticum, and cavum

acustico-jugulare
cavum tympanicum (Ogushi, 1911) = cavum tympani
choana (Siebenrock, 1897; Kesteven, 1910; Ogushi, 1911; Nick, 1912) = apertura

narium interna
columella (Siebenrock, 1897) = columella auris
columella (Ogushi, 1911) = os epipterygoideum
complementare (Ogushi, 1911) = os coronoideum
crista basioccipitalis (Nick, 1912) = crista dorsalis basioccipitalis
crista occipitalis (Kesteven, 1910; Ogushi, 1911) = crista supra-occipitalis
crista praetemporalis (Siebenrock, 1897) = processus trochlearis
dorsum ephippii (Siebenrock, 1897) = dorsum sellae
fenestra rotunda (Kesteven, 1910) = fenestra perilymphatica
foramellum anterius pro nervo glossopharyngeo (Ogushi, 1911) = foramen internum

nervi glossopharyngei
foramellum posterius pro nervo glossopharyngeo (Ogushi, 1911) = foramen externum

nervi glossopharyngei
foramen acusticum (Nick, 1912) = foramen nervi acustici
foramen acusticum anteriore principale (Ogushi, 1911) = foramen nervi acustici

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 103

foramen alveolare superius anterius (Ogushi, 1911) = foramen alveolare superius
foramen arteriae facialis (Nick, 1912) = foramen stapedio-temporale
foramen carotico-temporale (Siebenrock, 1897) = foramen stapedio-temporale
foramen caroticum (Kesteven, 1910) = foramen anterior canalis carotici interni
foramen caroticum externum (Siebenrock, 1897) = aditus canalis stapedio-temporalis
foramen caroticum internum (Siebenrock, 1897) = foramina anterior et posterior

canalis carotici interni
foramen carotidis internae (Nick, 1897) = foramen anterior canalis carotici interni
foramen cochleae (Siebenrock, 1897) = fenestra perilymphatica
foramen cribrosum anterius (Kesteven, 1910) = foramen nervi acustici
foramen externum nervi facialis (Nick, 1912) = foramen nervi facialis (in part)
foramen externum nervi hypoglossi (Nick, 1912) = foramen nervi hypoglossi (in

part)
foramen incisivum (Siebenrock, 1897; Ogushi, 1911; Nick, 1912) = foramen

praepalatinum
foramen internum nervi facialis (Nick, 1912) = foramen nervi facialis (in part)
foramen internum nervi hypoglossi (Nick, 1912) = foramen nervi hypoglossi (in part)
foramen jugulare internum (Siebenrock, 1897) = foramen cavernosum
foramen jugulare internum (Ogushi, 1911) = foramen jugulare anterius
foramen lacerum (Siebenrock, 1897; Ogushi, 1911) r= fenestra postotica
foramen lachrymale (Kesteven, 1910) = foramen orbito-nasale
foramen narium externum (Ogushi. 1911) apertura narium externa
foramen nasopalatinum (Nick, 1912) = foramen orbito-nasale
foramen nervi abducentis (Nick, 1912) = canalis nervi abducentis
foramen nervi glossopharyngei (Siebenrock, 1897) = foramen externum nervi glosso-

pharyngei
foramen nervi vidiani, ramus facialis (Siebenrock, 1897) = foramen pro ramo nervi

vidiani
foramen occipitale (Siebenrock, 1897) = foramen magnum
foramen occipitale magnum (Ogushi, 1911) = foramen magnum
foramen ovale (Kesteven, 1910) = foramen nervi trigemini
foramen ovale (Ogushi, 1911) = fenestra ovalis

foramen palatino-nasale (Siebenrock, 1897) = foramen orbito-nasale
foramen perilymphaticum (Nick, 1912) := fenestra perilymphatica
foramen posterior canalis carotici (Ogushi, 1911) = foramen posterior canalis

carotici interni
foramen pro nervo glossopharyngeo (Siebenrock, 1897) = foramen internum nervi

glossopharyngei
foramen pro nervo hypoglosso (Ogushi, 1911) = foramen nervi hypoglossi
foramen sphenoidale (Siebenrock, 1897; Ogushi, 1911; Nick, 1912) = foramen nervi

trigemini
foramen temporale (Kesteven, 1910) = foramen stapedio-temporale
foramen vago-accessorium (Kesteven, 1910) = foramen jugulare anterius
foramen vestibuli (Siebenrock, 1897) = fenestra ovalis
fossa hypophyseos (Siebenrock, 1897; Nick, 1912) = sella turcica
fossa jugularis (Ogushi, 1911) = cavum acustico-jugulare (in part)
fovea articularis (Siebenrock, 1897) = area articularis mandibularis

104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

fovea major (Nick, 1912) = recessus labyrinthiciis prooticus

goniale (Nick, 1912) = os praearticulare

hiatus posterior canalis carotici (Kesteven, 1910) = foramen posterior canalis

carotici interni
incisura columellae (Siebenrock, 1897) = incisura columellae auris
incisura ductus cochlearis (Ogushi, 1911) = foramen aquaducti vestibuli
incisura jugularis posterior (Siebenrock, 1897; Nick, 1912) = foramen jugulare

posterius
intermaxillare (Ogushi, 1911; Nick, 1912) = os praemaxillare
lamina alisphenoidea (Kesteven, 1910) = processus inferior parietalis
meatus auditorius (Siebenrock, 1897) = fossa acustico -facialis
meatus auditorius internus (Ogushi, 1911) = fossa acustico-facialis
operculare (Siebenrock, 1897; Ogushi, 1911) = os praearticulare
operculare (Nick, 1912) = os spleniale

os occipitale basilare (Kesteven, 1910) = os basioccipitale
os occipitale laterale (Kesteven, 1910) = os exoecipitale
os occipitale superius (Kesteven, 1910) = os supraoccipitale
os postfrontale (Kesteven, 1910) = os postorbitale

ostium superius canalis inframaxillaris (Ogushi, 1911) ==■ fossa meckelii
otosphenoideum (Siebenrock, 1897) = os prooticum
paraquadratum (Siebenrock, 1897) = os quadrato-jugale
paraoccipitale (Siebenrock, 1897) = os opisthoticum
pars sellae (Kesteven, 1910) = sella turcica

pediculus ossis quadrati (Kesteven, 1910) = processus epipterygoideus
pleuroccipitale (Siebenrock, 1897) = os exoecipitale

postfrontale (Siebenrock, 1897; Ogushi, 1911; Nick, 1912) = os postorbitale
praeoperculare (Siebenrock, 1897) = os spleniale

processus clinoideus anterior (Ogushi, 1911) = rostrum basisphenoidale
processus clinoideus posterior (Ogushi, 1911) = processus clinoideus
processus condyloideus occipitalis (Ogushi, 1911) = condylus occipitalis
processus ectopterygoideus (Siebenrock, 1897; Nick, 1912) = processus pterygoideus

externus
processus epipterygoideus (Kesteven, 1910) = crista pterygoidea
processus labyrinthicus (Ogushi, 1911) = processus interfenestralis
processus paroccipitalis (Kesteven, 1910) =: tuberculum basioccipitale
processus posterior basioccipitalis (Ogushi, 1911) = tuberculum basioccipitale
processus pterygoideus (Nick, 1912) = processus epipterygoideus
processus trabecular inferiores (Siebenrock, 1897) = rostrum basisphenoidale
ramus suspensorius (Kesteven, 1910) = processus articularis
recessus ampullae posterioris (Nick, 1912) = recessus labyrinthicus opisthotieus
recessus cavum tympani (Siebenrock, 1897) = cavum acustico-jugulare
recessus posterior ampullaris (Ogushi, 1911) = recessus labyrinthicus opisthotieus
recessus pro sinu superiore utriculi (Nick, 1912) = recessus labyrinthicus supra-

occipitalis
recessus superior (Ogushi, 1911) = recessus labyrinthicus supraoccipitalis
recessus vestibularis anterior (Ogushi, 1911) = recessus labyrinthicus prooticus
recessus vestibularis opisthotieus (Kesteven, 1910) = recessus labyrinthicus opisthoti-
eus

PARSONS AND WILLIAMS: JURASSIC TURTLE SKILLS 105

recessus vestibularis prooticus (Kesteveu, 1910) = recessus labyrinthicus prooticus

rima nasalis (Ogushi, 1911) = fissura ethmoidalis

rostrum ossis basisphenoidei (Kesteven, 1910) = rostrum basisphenoidale

sinus mastoideus (Ogushi, 1911) = antrum postoticum

sulcus columellae amis (Kesteven, 1910) = incisura columellae amis

sulcus ethmoidalis (Kesteven, 1910) = sulcus vomeri

sulcus inframaxillaris (Ogushi, 1911) = sulcus cartilaginis meckelii

sulcus jugularis (Kesteven, 1910) = sulcus cavernosus

sulcus pro nervo vago et accessorio (Ogushi, 1911) = foramen jugulare posterius

sulcus vestibularis epioticus (Kesteven, 1910) = recessus labyrinthicus supraoccipitalis

supraangulare (Siebenrock, 1897; Nick, 1912) = os surangulare

suprangulare (Ogushi, 1911) = os surangulare

trabeculae (Nick, 1912) = rostrum basisphenoidale

trochlea prootica (Ogushi, 1911) = processus trochlearis

tuberculum ovoideum (Kesteven, 1910) = crista dorsalis basioecipitalis

vestibulum (Siebenrock, 1897; Nick, 1912) = cavum labyrinthicum

ACKNOWLEDGMENTS

This work has been an outgrowth of studies undertaken during
tenure of a Guggenheim Fellowship by the junior author in the years
1952-1953.

The Geology Department of the British Museum (Natural History),
their preparation staff and the museum's photographic department pro-
vided every assistance during the visit of the junior author. Dr. W. E.
Swinton was especially helpful and encouraging. Dr. C. K. van Regteren
Altena of the Teyler Museum permitted the removal by acetic acid of
the limestone matrix that initially concealed the Solenhofen skull. The
patience of these authorities during the very lengthy period of the loan
of this material is very gratefully acknowledged.

The Staatliches Museum fiir Naturkunde, Stuttgart, graciously pro-
vided opportunities for the examination of the superb skeletons of
Proganochelys and very generously furnished photographs which are
published here. Dr. K. D. Adam in particular gave of bis time and
energy during the junior author's study of this material.

Miss Patricia Washer, Mrs. Dorothy Marsh, Miss Margaret Estey,
and Dr. Margaret C. Parsons prepared the drawings.

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1929. Studies in chelonian osteology. Part II. The morphological relationships
between the girdles, ribs and carapace. Ann. New York Acad. Sci., vol. 31,
pp. 81-120.
Schumacher, G. H.

1954. Beitrage zur Kiefermuskulatur der Schildkroten. 1. Mitteilung. Bail

des M. adductor mandibularis miter spezieller Beriicksiehtigung des M.

pterygoideus bei Chelone, Caretta, Podocnemis, Pelusios und Testudo

elephantopus. Wissensch. Zeitschr. Univ. Greif swald, Jahrg. 3, pp. 149-210.

Seeley, H. G.

1892. On a new reptile from Welte Vreden (Beaufort West), Eunotosaurus
africanus (Seeley). Quart. J. Geol. Soc. London, vol. 48, pp. 583-585.
Seydel, O.

1896. Uber die Nasenhohle und das Jacobson' sche Organ der Land- und
Sumpfschildkroten. Festschr. 70 Geburtstage Gegenbaur, vol. 2, pp.

385-486.

PARSONS AND WILLIAMS: JURASSIC TURTLE SKULLS 107

SlEBENROCK, F.

1897. Das Kopfskelet der Schildkvoten. Sitzungsb. Kais. Akad. Wissenseh.

(Wien), Math.-Naturw. Kl., vol. 106, Abth. 1, pp. 245-328.
Walker, W. F., Jr.

1947. The development of the shoulder region of the turtle, Chrysemys picta

marginata, with special reference to the primary musculature. J. Morph.,

vol. 80, pp. 195-249.
Watson-, D. M. S.

1914. Eunotosaurus africanus Seeley, and the ancestry of the Chelonia. Proc.

Zool. Soc. London, 1914, pp. 1011-1020.
Williams, E. E.

1954. A key and description of the living- species of the genus Podocnemis

{sensu Boulenger ) (Testudines, Pelomedusidae ). Bull. Mus. Comp. Zool.,

Harvard Coll., vol. Ill, no. 8, pp. 279-295.
Zangerl, E.

1939. The homology of the shell elements in turtles. J. Morph., vol. 65, pp.

383-409.
1953. The vertebrate fauna of the Selma Formation of Alabama. Part III.

The turtles of the family Protostegidae. Part IV. The turtles of the

family Toxochelyidae. Fieldiana : Geol. Mem., vol. 3, nos. 3 and 4, pp.

57-277.

PLATES

1. Portland skull. British Museum R2914. Palatal view.

(Courtesy of the British Museum)

2. Portland skull. British Museum R2914. Left lateral view.

(Courtesy of the British Museum)

3. Upper: Portland skull. British Museum R3163. Anterior view of
skull fragment. (See Text-fig. 8).
Lower : Portland mandible. British Museum R2914. Dorsal view.

(Courtesy of the British Museum)

4. Solenhofen skull. Teyler Museum.

Upper : dorsal view. Lower : palatal view.

5. Proganochelys quenstedti, Stuttgart skull. Upper: lateral view.
Lower: palatal view.

(Courtesy of the Stuttgart Museum)

6. Proganochelys quenstedti, Stuttgart skull. Upper: anterior view.
Lower : posterior view.

(Courtesy of the Stuttgart Museum)

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 125 No. 4

ON THE SPECIES OF PSEUDIDAE (AMPHIBIA, ANIIRA)

By J. M. Gallardo
Museo Argentine) de Ciencias Naturales, Buenos Aires.

With One Plate

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Bulletin of the Museum of Comparative Zoology

AT HAEVARD COLLEGE

Vol. 125, No. 4

ON THE SPECIES OF PSEUDIDAE (AMPHIBIA, ANURA)

By J. M. Gallardo
Museo Argentino de Cieneias Naturales, Buenos Aires.

With One Plate

CAMBRIDGE, MASS., U.S.A.
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Xo A --On tin Species of Pseudidae {Amphibia. Anura)

By J. M. Gallardo

INTRODUCTION

A very substantial contribution toAvards an understanding of
the systematies of the frog's of the genera Pseuelis and Lysapsus
was achieved by Savage and Carvalho (1953). A few points,
however, appear to require modification, among them the status
of Pseudis mantidaetylus Cope (thought by Savage and Carval-
ho to be a synonym of P. paradoxus) .

A preliminary examination of specimens of Pseudis mantidae-
tylus in the collection of the Seccion Herpetologia del Museo
Argentino de Ciencias Naturales, of the Facultad de Ciencias
Exactas y Naturales de la Universidad de Buenos Aires and of
the Colegio Nacional de Buenos Aires (Argentina), as well as of
others collected by myself in Argentina, Provincia de Buenos
Aires (Paso de la Noria, San Miguel, Bella Vista, Santo Do-
mingo ) , impressed upon me the constancy of the shape and posi-
tion of the dark and light bands of the ventral aspect of the
thighs. In contrast, specimens of Pseudis paradoxus from Ro-
sario, Santa Fe and Corrientes, showed a quite different thigh
pattern. The specimens of P. paradoxus from other areas in
the Museum of Comparative Zoology (MCZ), United States
National Museum (USNM), the American Museum of Natural
History (AMNH), Chicago Natural History Museum (CNHM),
and Carnegie Museum (CM) collections, which I was able to
examine in 1959-60, showed still other patterns of thigh colora-
tion. Such facts as these led me to regard P. mantidaetylus as a
distinct species and to recognize several subspecies within P.
paradoxus as well. A study of Lysapsus limellus showed the ex-
istence of races in this form also.

My previous studies on Bufo granulosus and its subspecies
have convinced me that there has been recognition of too many
species and subspecies without proper reference to the ecological
and physiographical features of the environment. I have attempt-
ed to study the present species in relation to natural geograph-
ical units, such as the hydrographical systems of South Ameri-
ca. Such a procedure is particularly useful for the Pseudidae,
which are exceedingly aquatic. The species I know best is P.
mantidaetylus. I am familiar with it in the wild and I have
never seen it come out of the water voluntarily ; instead, it stays
always in pools and slow creeks, covered with floating vegetation
(Lemna sp. and Jussiaea sp.). Fernandez and Fernandez (1921:

112 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

139) also report that P. mantidactylus in captivity does not
leave the water, remaining in small muddy puddles. Possibly in-
dividuals move from pool to pool following heavy rains ; ordin-
arily, however, the several populations very definitely tend to
remain each within a given hydrographical basin. This is very
apparent in the distributional patterns ; of the six subspecies of
Pseudis paradoxus which I describe below, one is restricted to the
Guianas, another to the Orinoco system, others to the Sao Fran-
cisco, Arassuahy and Beni rivers and a last one to the Kio de la
Plata basin. It is not improbable that further subspecies may
be recognized in the intervening regions.

PSEUDIS PARADOXUS (LINNAEUS)

Description of the species as a whole. Snout more or less sharp,
canthus rostralis indistinct. Loreal region oblique. Tympanum
distinct. Vomerine teeth between the choanae large, forming two
straight bars, little separated. Tongue rounded, with a small
notch. Male with a single internal vocal sac. Small granulations
on the dorsum, extending on to the hind limbs. In the pectoral
region two sharp angular marks, corresponding to the scars pro-
duced in metamorphosis by the eruption of the fore limbs (a
character persistent in the adult). Fore limbs relatively weak;
fingers long, the pollex opposable ; all fingers with cutaneous frin-
ges, and with basal interdigital membranes between the second,
third and fourth fingers ; a long internal palmar tubercle and a
small external one ; simple subarticular tubercles. Proximal part
of the arm wholly included in skin of body. Hind limbs strong,
thigh and leg broad. Tarsal fold present ; internal metatarsal
tubercle more or less developed (varying according to sub-
species) ; no external metatarsal tubercle, cutaneous fringe on
the outermost and innermost toes ; interdigital membrane well
developed, inserting between the metatarsals and extending to
the terminal half of the dilatations ; subarticular tubercles sim-
ple. Dorsal coloration intermediate between greenish and brown,
with darker spots. Venter with more or less abundant brown
spotting. Four longitudinal dark lines and alternate light spaces
in the ventral aspect of the thigh (the shape and disposition of
these dark lines vary in the different subspecies ; in the descrip-
tions I shall number them from 1 to 4, starting from the external
edge). A long light spot over the cloaca and one or two below it,
prolonged by a row of light spots.

G-ALLARDO : SPECIES OF PSEUDIDAE

11:;

Skeletal characters. Skull with swollen otic region; occipital
condyles widely separated. Correlated with this, the atlas with
two anterolateral prominences to receive the condyles which are
far apart, the space between the prominences concave (i.e. "odon-
toid process" absent). [A similar atlas exists in some Leptodac-
tylidae (Leptodactylus, Eleutherodactylus) , Hylidae (Hyla),
some Ranidae (Arthrolcpticles, Oeidozyga) and Phrynomeridae

Fig. 1. Atlas and occiput of Bufo spinulosus ( left) compared with atlas
and occiput of Pseudis paradoxus (right).

(Phrynomcrus). In other Leptodactylidae (Batrachophrynus,
Telmatobius, Ceratophrys, Calyptocephala), Pelobatidae($cap/w-
opus), Pelodytidae (Pclodytes), some Ranidae (Astylosternus,
Rana) and Bufonidae (Bufo), there are no lateral prominences;
instead the median portion of the articular surface is convex (i.e.
"odontoid process" present) and the occipital condyles are closer
to each other. This characteristic of the Pseudis atlas makes it
inadvisable to group Batrachophrynus, Telmatobius, Ceratophrys
and Calyptocephala with Pseudis in a subfamily Pseudinae as
Noble suggested. In addition, however, the presence of an acces-
sory phalanx in all digits excludes from the Pseudinae not only
the genera just mentioned but also other genera such as Eleu-
therodactylus which Noble also wished to place in the subfamily,
and which do show the Pseudis type of atlas.] Transverse
processes of the vertebrae, including the second and sacral, ap-
proximately cylindrical ; at the proximal end of the urostyle there
is a neural arch as in a 10th vertebra (cf. some skeletons of
Batrachophrynus macrostemum) • close to this there is a 10th
pair of the calcarean sacs. Pectoral girdle arciferal, omosternum
plus a xiphisternum as a short, notched, cartilaginous plate.
Terminal phalanges long, reaching the tips of the digits, i.e. discs
not present.

114

BULLETIN' : MUSEUM OF COMPARATIVE ZOOLOGY

Fig. 2. Pectoral girdle of Lysapsus mantidactylus.

Fig. 3. Outline of tip of toe in relation to terminal phalanx. Lysapsus
mantidactylus, left, dorsal view; right, lateral view.

SUBSPECIES OP PSEPDIS PARADOXUS

To differentiate the subspecies of Pseudis paradoxus, I use the
following morphological characteristics : size of vomerine teeth,
size of tympanum, whether the supratympanic fold is more or
less visible, the shape of the internal metatarsal tubercle ; and the
following color characters: number of dark bands on the back,
ventral spots on throat, breast and abdomen, dark lines and light
spaces on the ventral thigh, one or two light spots below cloaca,
and coloration of the ventral tibia.

Pseudis paradoxus paradoxus (L.)

1 758 Bana paradoxa Linnaeus, p. 212.

1830 Pseudis paradoxa Wagler, p. 203; Gunther, 1858, p. 5;

Boulenger, 1882, p. 186.
Description. Adult male MCZ 12136, Demerara, British
(luiana. The longitudinal diameter of the tympanum is equal to

(IALLARDO: SPECIES OF PSEUDIDAE 115

eye diameter and also to the distance from eye to nostril. Supra-
tympanic fold not distinct. Vomerine teeth large, with little sep-
aration. Metatarsal tubercle conical, oblique, not hooked. Back
with four longitudinal dark lines, more or less interrupted. Dark
spots on the dorsal aspect of limbs -- longitudinal on the anterior
limbs and transverse or oblique on the posterior limbs; external
edge of the tibia with a dark longitudinal line ; internal edge of
tarsus dark. Dark spots on the interdigital membrane. Brown
irregular spots on the throat and the ventral aspect of the fore
limbs. On tbe pectoral region a dark comma-shaped line on each
side at the pectoral angular scars; another pair of more lateral
spots. Abdomen with scattered small round spots. Ventral sur-
face of the thigh with longitudinal dark lines not very wide,
sometimes broken up and with angular projections; the space
between line 1 and the external edge is light and with irregular
long spots ; space between lines 1 and 2, light, immaculate, without
invading angular projections from the bounding lines ; space be-
tween 2 and 3 with such angular projections ; lines 3 and 4 fre-
quently joined towards the middle of the femur ; space between
line 4 and internal edge with angular projections that form light
rectangles. Two light long spots below the cloaca. Ventral as-
pect of tibia with abundant short irregular lines.

Dimensions : Head and body 56 mm. Head length 18 mm. Head
width 19 mm. Head height 8 mm. Eye 6 mm. Interorbital space
6 mm. Elbow to third finger 30 mm. Femur length 30 mm. Tibia
length 32 mm. Tibia width 11 mm. Heel to fourth toe 45 mm. Foot
length 32 mm. Adult female, same origin, MCZ 12135, head and
body 54 mm.

Distribution : This subspecies is restricted to the British and
Dutch Guianas.

Material studied: MCZ 12135-6 (2 specimens) British Guiana,
Demerara, coast lands, E. Smith; MCZ 2775 (1 specimen) British
Guiana, Demerara, C. W. Beebe, 1909 ; AMNH 13566-8 (3 speci-
mens) British Guiana, Demerara River, Alayma, J. Rodway;
MCZ 3901-2 (2 specimens) British Guiana, trenches near George-
town, J. Rodway, 1914; AMNH 1305 (1 specimen) British
Guiana, J. Rodway, 1912; AMNH 21423, 39588, 39638-40 (5 spe-
cimens) British Guiana, Georgetown ; AMNH 49259 (1 specimen)
British Guiana, Essequibo River, R. Snediger, V-31-1937 ; AMNH
5103 (1 specimen) Dutch Guiana, Paramaribo, J. A. Samuels;
AMNH 24013 (1 specimen) Guiana.

116 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

PSEUDIS PARADOXUS CARIBENSIS Sllbsp. 110V.

1933 Pscudis paradoxa Parker, p. 10; 1934, p. 123; Gans,
1956, p. 2 (part) ; Kenny, 1956, p. 23.

1942 Pseudis paradoxis Ditmars, p. 51.

For diagnostic features see Table 1 .

Description. Type, adult female MCZ 19890, Mayaro Bay,
Trinidad, B. W. I. Tympanum rather smaller than the eye and
smaller than the distance between eye and nostril. Oblique fold
above the tympanum indistinct, but marked by a dark line. Vo-
merine teeth large, with little separation. Metatarsal tubercle
with narrow base, forming a small hook. Back with six longitu-
dinal dark lines more or less discontinuous. Dorsal surface of
limbs and interdigital membrane as in the preceding subspecies.
Throat, pectoral and abdominal regions with dark rounded spots
(more abundant on the throat and breast). Ventral surface of
thigh with thinner lines than in /'. p. paradoxus; space between
line 1 and external edge, with elongated punctations and spots;
space between lines 1 and 2 light ; space between lines 2 and 3 with
abundant elongate spots; space between 3 and 4 light (the fourth
line wider) ; between line 4 and internal edge abundant, elongate,
irregular spots. One light, long, spot below the cloaca. Tibia
ventrally with abundant short, irregular, lines. (The ventral
coloration of the hind limbs is also perfectly visible in four-legged
tadpoles.)

Dimensions : Head and body 64 mm. Head length 22 mm. Head
width 25 mm. Head height 11 mm. Eye 6 mm. Snout height 4
mm. Interorbital space 7 mm. Elbow to third finger 33 mm.
Femur length 39 mm. Tibia length 33 mm. Tibia width 11 mm.
Heel to fourth toe 50 mm. Foot 36 mm. Paratype, adult male,
CM 33787, head and body 60 mm.

Distribution: Nearly all the material that has been studied by
me comes from the same locality, Mayaro, S.E. Trinidad Island.
(This locality has recently been described, and photographs of it
published by dans, 1956.) Additional Trinidad localities have
been reported by Kenny (1956) : Plum Mitan ricefields near
Biche (tadpoles) ; Icacos swamp near Cedros (adults) ; Apex oil-
fields at Fyzabad (males singing). This subspecies, however, seems
to exist also in Venezuela. At least the specimen of Pseudis para-
eloxus figured by Lutz (1927, Plate II, figs. 18-19), from Maracay,
Venezuela, has a ventral pattern similar to the Trinidad speci-
mens. I have seen also one specimen from Hato Gabinero, Vene-
zuela, that in general has the characteristics of this subspecies. In

GALLARDO: SPECIES OF PSEUDIDAE 117

Ibis specimen the ventral coloration of the thighs is nearly the same
as typical P. p. caribensis, with spots in the space between line 1
and external edge; some spots (but few and small) in the distal
portion of the space between 2 and 3; short and irregular lines
between the knee and the union of 3 and 4 ; moreover the comma-
shaped spots on the pectoral region are lacking and the metatarsal
1 nbercle is hook shaped. Other characteristics are close to those of
P. p. paradoxus, as is the presence of four interrupted longitu-
dinal dark dorsal bands; it is, however, different from both sub-
species by having the ventral surface of the tibia without spots.

Material studied : MCZ 19890-1 ; 19893 (3 specimens) Trinidad,
Mayaro Bay, N. A. Weber, XIi-1934; MCZ 19892 (1 skeleton)
Trinidad, Mayaro Bay; MCZ 26157 (1 specimen) Trinidad, St.
Bernard Estate. Mayaro, M. G. Kugler, 1945; CXHM 49705-6 (2
specimens) Trinidad, Plaisance Mayaro, Stander, 1947 ; CM 33787
(1 specimen) Trinidad, Pond at Mayaro, C. Gans, IX-22-1953 ;
LFSNM 137347 (1 specimen) Venezuela, Cojedes, Hato Gabinero
near San Carlos, Perez, IY-8-1955.

PSEUDIS PARADOXUS BOLBODACTYLUS A. Lutz

1925 Pseuelis bolbodactyla A. Lutz, p. 138; Cochran, 1955,
p. 312, fig. 20, PI. 32, figs. G, H.

Description. Female, MCZ 25729, Lagoa do Curralinho, Las-
sance, Minas Gerais, Brasil. Tympanum rather smaller than the
eye and smaller than the distance from eye to nostril. Supratym-
panic fold not very evident. Vomerine teeth more reduced than
in the two preceding subspecies and separated on the mid line.
Metatarsal tubercle more or less conical, not forming a hook. An
interocular spot produced posteriorly on the two sides in two
broad arms with an indentation between, thence continued on the
back by two longitudinal irregular lines. Dorsal aspect of limbs
with spots, internal edge of the femur with large light-edged
ocelli. Interdigital membrane of the foot with spots barely evi-
dent. Throat, pectoral and abdominal regions with dark rounded
spots, more abundant on the breast. Ventral region of thigh with
relatively thick lines, but several of them interrupted, space be-
tween line 1 (which is rather curved) and the external edge light ;
proximally between lines 1 and 2 an elongate spot (line 2 starts
with two spots) ; space between lines 2 and 3 light (line 3 com-
posed of elongate spots) ; space between 3 and 4 light (line 4 is
formed by round spots ) ; space between 4 and internal edge light.
One light long spot below the cloaca. Ventral aspect of tibia with
a few dark spots.

118 BULLETIN: MUSEUM OE COMPARATIVE ZOOLOGY

Dimensions : Head and body 45 mm. Head length 16 mm. Head
width 17 mm. Head height 8 mm. Eye 4 mm. Interorbital space
3 mm. Elbow to third finger 22 mm. Femur length 25 mm. Tibia
length 24 mm. Tibia width 9 mm. Heel to fourth toe 35 mm. Foot
27 mm.

Variation : In general the lines on the thigh are not so much
interrupted as in the specimen described, especially lines 3 and 4,
which are usually continuous or have one spot at the beginning or
at the end. More constant is the short line in the space between
1 and 2 (sometimes united to line 2 and forming an angle) and
the spot at the beginning of line 2. This subspecies has many
similarities with P. p. platensis, but it is different in having line
1 thinner, no angular projections from any of the lines and no
fusion of lines 3 and 4.

Distribution: Confined to the Sao Francisco River in the State
of Minas Gerais, Brasil. Cochran (1955 : 215) cites material from
various localities in this state and from lta in the state of Espirito
Santo, though it would be interesting to check whether the present
subspecies or another one exists in the latter locality.

Material studied: MCZ 25729 (1 specimen) Brasil, Minas
Gerais, Lassance, Lagoa do Curralinho, Cochran and Venancio ;
USNM 98534 (1 specimen) Brasil, Pirapora, 111-23-1935 ; USNM
98173, 175-79, 181-88, 190-95, 197-99, 200, 202 (25 specimens)
Brasil, Lassance, Lagoa do Curralinho, Cochran and Venancio,
111-22-1935; USNM 97022 (1 cotype) Brasil, Minas Gerais, Las-
sance, Lagoa de Genipapo.

Pseudis paradoxus fuscus Garinan

1883 Pseudis fusea Garman, p. 47; Cochran, 1955, p. 216, PI.
32, figs. I-K.

Description. Cotype MCZ 1872, Rio Arassuahy, Minas Gerais,
Brasil. Tympanum approximately the same size as the eye and
as the distance from eye to nostril. Supratympanic fold present.
Snout relatively short (a variable character in P. p. paradoxus) .
Vomerine teeth relatively reduced and separated in the middle.
The metatarsal tubercle not forming a hook. Dorsal color pattern
rather indistinct ; ground color of cotype brown, perhaps due to
poor preservation ; ventral parts light, rather spotted towards the
throat. In the cotype it is practically impossible to see the colora-
tion of the ventral aspect of the femur, though it would seem that
the space between line 1 and the external edge has abundant dark

GALLARDO : SPECIES OF l'SEI'DIDAE

11!)

spots. Space between line 4 and the internal edge, with large ir-
regular light spots. A light line above the cloaca; another light
line below it, nearly divided into two spots and continued by
other light spots. Ventral aspect of tibia with abundant irregular
dark spots.

Fig. 4: Ventral aspects of thighs in Pseudis and Lysapsus. Upper row:
left, P. p. paradoxus MCZ 12135 ; center, P. p. caribensis MCZ 19890 ; right,
P. p. bolhodactylus MCZ 25729. Lower row: left, P. p. platensis MCZ
2-4808; center, P. p. ocddentalis MCZ 299(il ; right, Lysapsus mantidactylus
MCZ 28730.

Dimensions: Head and body 51 mm. Head length 16 nun. Head
width 1!) mm. Head height 8 mm. Eye 6 mm. Interorbital space
4 mm. Elbow to third finger 26 mm. Femur 20 mm. Tibia length
28 mm. Tibia width 9 mm. Heel to fourth toe 36 mm. Foot 26 mm.
Another eotype, head and body 41 mm.

Distrib ution : No specimens have been reported since Garman 's,
and the only known locality is the Arassuahy River in the State
of Minns Gerais. This locality is very close to that of P. p. bol-
bodactylus, but separated by the Serra do Espinhaco which
divides the Sao Francisco basin from that of the Jequitinhonha

120 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

basin, to which the Arassuahy River belongs. Partly on the basis
of this geographical separation (cf. Cochran 1955: 217) and in
spite of the absence of new material, I am disposed to retain this
form as distinct from P. p. bolbodactylus.

Material studied: MCZ 1872 (2 specimens) cotypes, Brasil,
Minas Gerais, Rio Arassuahy, Hartt and Copeland. Thayer Ex-
pedition, 1864-7.

PSEUDIS PARADOXUS PLATENSIS SUbsp. 110V.

1894 Pseudis paradoxa Boulenger, p. 347 ; Miranda Ribeiro,
1926, pi. II, figs. 1-la; Muller and Hellmich, 1936, p. 27,
fig. 9; Freiberg, 1942, p. 228 (part); Travassos and
Freitas, 1942, p. 284; Vellard, 1948, p. 173; Cei, 1956,
pi. V, figs. 36-37; Gans, 1960, p. 303 (part).

For diagnostic features see Table 1.

Description. Type, adult male, MCZ 24808, Colonia Xueva
Italia, Dept. Villeta, Paraguay. Tympanum approximately the
same size as the eye and as the distance from eye to nostril. Prac-
tically no supratympanic fold. Vomerine teeth large, with little
separation. Metatarsal tubercle rather prominent, but without
hook. On the back two large dark spots shaped like commas, ex-
tending from the interorbital space to the scapular region; two
interrupted dorsal dark lines. Dorsal aspect of the limbs and
interdigital membrane of the foot with dark spots. Throat, pec-
toral and abdominal regions with more or fewer round dark
spots. ATentral aspect of the thigh with dark thick lines; space
between line 1 and external edge without spots; space between
line 1 and line 2 with an extensive proximal spot; light space
between line 2 and line 3 with the projections from the lines only
slightly indicated ; space between line 3 and line 4 light ; between
line 4 and the internal edge, light rectangles. Two light long spots
below the cloaca. Ventral aspect of tibia with extended dark
spots interrupted and irregular.

Dimensions : Head and body 50 mm. Head length 15 mm. Head
width 16 mm. Head height 7 mm. Eye 5 mm. Interorbital space
3 mm. Elbow to third finger 24 mm. Femur length 28 mm. Tibia
length 25 mm. Tibia width 8 mm. Heel to fourth toe 36 mm. Foot
25 mm. Paratype MCZ 24809. from the same locality, head and
body 42 mm.

Distribution : This subspecies is found from the Upper Para-
guay River to Rosario (Santa Fe) on the banks of the Parana

GALLAKDO : SPECIES OF PSEUDIDAE

121

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122 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

River. In Brasi] it has been collected in Mato Grosso: S. Luiz de
Caceres (the northernmost locality known at present) and near
Bodoquena ; in Bolivia, San Fermin at 100 km NW of Puerto
Suarez and in nearby El Carmen; in Paraguay, near Asuncion.
Fortin Esteros and Nueva Italia ; in Argentina in the Provinces
of Misiones, Corrientes (Ituzaingo, Isla Apipe) and Santa Fe
(La Geraldina, Santa Fe, Rosario).

Material studied: MCZ 24808-9 (2 specimens) Paraguay, Dept.
Villeta, Colonia Nueva Italia, P. Willim, XII-6-1943; CNHM
42313-4 (2 specimens) Paraguay, Colonia Nueva Italia, P. Willim ;
AMNII 50657 (1 specimen) Paraguay, Villeta, Colonia Nueva
Italia, P. Willim; MCZ 29958 (1 specimen) Bolivia. El Carmen,
C. Cans, 11-26-1954.

PSEUDIS PARADOXUS OCCIDENTALIS Sllbsp. 110 V.

1956 Pseudis paradoxa dans. p. 2 (part) ; (ians. 1960, p. 303
(part).

For diagnostic features see Table 1 .

Description. Type, adult male, MCZ 29961, El Pailon, Bolivia.
Tympanum slightly smaller than the eye and smaller than the
distance from eye to nostril. The skin of the supratympanic fold
covers part of the tympanic membrane. Loreal region slightly
concave. Vomerine teeth large, very slightly separated. Meta-
tarsal tubercle rather prominent, but without hook. Back brown
with four black longitudinal lines which are very irregular and
have lateral projections, forming dark edged circles towards the
rear. Dorsal aspect of the extremities with dark spots. Internal
edge of the tarsus dark. Interdigital membrane of the foot with
large dark spots. Sides of trunk with light spots. Ventral aspect
of thigh with the four dark lines very blurred (sometimes lines
1 and 2 are missing) ; lines 3 and 4 are more distinct and thicker;
the spaces among the lines are light. One light long spot below
the cloaca. The fourth ventral line of thigh follows the internal
edge. Tibia ventrally light. Throat, pectoral and abdominal re-
gions light and unspotted (in the male the throat slightly darker).

Dimensions: Head and body 51 mm. Head length 18 mm. Head
width 20 nun. Head height 9 mm. Eye 5 mm. Interorbital space
6 mm. Elbow to third finger 23 mm. Femur 25 mm. Tibia length
26 min. Tibia width 9 mm. Heel to fourth toe 35 mm. Foot 27 mm.
Paratypt MCZ 29959. adult female, the same locality, head and
bodv 72 mm.

GALLARDO : SPECIES OF PSEUDIDAE 123

Distribution : Though the material studied, with good locality,
comes from only one place in Bolivia, El Pailon, it is possible
that this susbspecies occupies the valleys of the Beni and its tribu-
taries. Recently, Shreve (1959) has described a new species of
Phyllomedusa (P. pailona) from the same locality, remarking
that Phyllomedusa sauvagii, its closest relative, was collected by
(rans in El Carmen: this distribution parallels that of Pseiidis
paradoxus occidentalis and P. p. platensis.

Material studied: MCZ 29959, 61 (2 specimens) Bolivia, El
Pailon, C. Gans, III-5-1954; USNM 94390 (1 specimen) South
America, J. II. Lunn, 1932.

INTERRELATIONSHIPS OF THE SUBSPECIES

P. p. paradoxus, P. p. earibensis and P. p. occidentalis of
northern and western South America seem to me more closely
related, while P. p. fuscus and P. p. bolbodactylus of southeast
Brasil form another group. P. p. platensis provides the connect-
ing link.

THE STATUS AND ALLOCATION OP
PSEUDIS MANTIDACTYLA COPE

This species must be considered distinct from Pseudis para-
doxus and more closely related to Lysapsus limellus. In general
it fits the definition of the genus Lysapsus given by Savage and
Carvalho (1953: 194). Thus it has terminal discs on the digits,
the interdigital webbing of the toes reaching to the base of the
discs: the vomerine teeth arranged obliquely, slightly towards the
rear of the choanae ; the pectoral girdle with a long and cartilagin-
ous omosternnm, of the same length as the epicoracoids. It differs
specifically from L. tinnitus in having the terminal phalanx of
the toes relatively not so short, although they do not reach the
extreme end of the digits on account of the presence of discs and
in the male with two vocal sacs. (The latter character also sepa-
rates mantidactylus from paradoxus.)

I propose therefore that Cope's species be called Lysapsus
mantidactylus (Cope). Parentheses are used since the generic
name employed by Cope (1862b: 352) in the original descrip-
tion was Pseudis and not Lysapsus, as wTrongly stated by Boulen-
ger (1882: 187), Freiberg (1942: 228), Savage and Carvalho
(1953: 193) and Cei (1956: 57).

124 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Lysapsus mantidactylus (Cope)

1862b Pseudis mantidactyla Cope, p. 352; Boulenger, 1882,
p. 187, 1883, p. 17 ; Boettger, 1892, p. 26 ; Berg, 1896, p.
162; Fernandez and Fernandez, 1921, p. 134; pi. Ill,
figs. 11-15; Nieden, 1923, p. 367 ; Miranda Ribeiro, 1926.
p. 25; Freiberg, 1912. p. 228; Gallardo, 1957, p. 354,
1958, pp. 291-298.
1953 Pseudis paradoxus Savage and Carvalho, p. 193 (part) ;

Cei, 1956, p. 57 (part).
Description. Adult female, MCZ 25730, Uruguay, Montevideo.
Tympanum equal to eye diameter and slightly larger than the
distance from eye to nostril. Head relatively shorter than in P.
paradoxus; practically no canthus rostralis. Vomerine teeth
short and separated, placed slightly to the rear of the choanae.
Small granulations on dorsum and hind limbs, forming lines on
the tibia. Leg (tibial region) thinner than in P. paradoxus. The
interdigital membrane only at the base of the fingers prolonged
onto the toes by digital fringes which reach the base of the ter-
minal discs. The metatarsal tubercle sharp and oblique, not
forming a hook. Tarsal fold not very prominent. One light
oblique band from the eye to the corner of the mouth. Back with
some dark spots. Sides of the trunk with a light longitudinal
band. A longitudinal dark band on the side of the fore limb.
Throat with some scattered spots, pectoral and abdominal regions
light. Ventral design of the femur: dark longitudinal lines, bet-
ter defined, in increasing order 1, 2, 3, 4; line 1 very blurred.
formed of long irregular spots ; spaces among the lines light. < )ne
light band over the cloaca and a fifth dark band at the level of the
cloaca; space between dark lines 4 and 5, light. Tibia ventrally
with infrequent spotting, more abundant on the edges.

Skeletal characters: Skeleton bluish green (in live animals
and freshly preserved specimens) ; Boulenger (1883: 17) makes
reference to this coloration. A green skeleton is also found in
other species of the genus, as in L. linn II us (Miranda Ribeiro,
1926: pi. II, fig. 2a), and according to Parker (1935: 510) in P.
laevis. Pectoral girdle arciferal ; coracoid and precoracoid ossified
and well developed; omosternum long and cartilaginous (same
length as the epicoracoids ) ; sternum and xiphisternum as a short
cartilaginous plate. The terminal phalanges are long, not reach-
ing the tips of the digits.

GALLARDO : SPECIES OF PSEUDIDAE 125

Dimensions: Head and body 35 mm. Head length 12 mm.
Head width 12 mm. Head height 4 mm. Eye 4 mm. Interorbital
space 2 mm. Elbow to third finger 18 mm. Femur length 20 mm.
Tibia length 19 mm. Tibia width 5 mm. Heel to fourth toe 27 mm.
Foot 19 mm.

Distribution : This species, first described by Cope from Buenos
Aires, was later reported by Boulenger and also by Boettger from
Rio Grande do Sul (Brasil) and Montevideo (Uruguay) ; by Berg
again from Buenos Aires, and Misiones (Argentina) ; and by Vel-
lard from Salta (Hickman) and Formosa (Puerto Dalmacia) in
Argentina. From the material I studied in Argentinian collections
and from my own collecting, I can say it is common in the neigh-
borhood of the city of Buenos Aires, extending to the N.E. of the
Buenos Aires Province and the lagunas of the Partido de General
Lavalle ; it can also be recorded for Santa Fe Province, where also
exist Pscudis paradoxus and Lysapsus limellus. Cei (1949: 539)
citer it for Corrientes.

Material studied: MCZ 25730 (1 specimen) Uruguay, Monte-
video; USNM 71124-6 (3 specimens) Uruguay, Puerto La
Paloma, W. L. Smith; USNM 70633-4 (2 specimens) Uruguay,
Montevideo ; USNM 65617-8 (2 specimens) Uruguay, Montevideo,
near Playa Ramirez, II. M. Smith, IX-30-1922 ; AMNII 11953 (1
specimen) Argentina, Prov. Buenos Aires, Punta Lara, Pedro
Serie. VI-15-1920; CNHM 9698 (1 specimen) Argentina, Prov.
Buenos Aires, La Plata, K. P. Schmidt, X-28-1926 ; USNM 22751-
2 (2 specimens) Argentina, Prov. Buenos Aires, La Plata; USNM
63506 (1 specimen) Argentina, Prov. Buenos Aires, Dolores, A.
Wetmore, X-1920 ; CNHM 83291 (1 specimen) Brasil, Rio Grande
do Sul ; CNHM 80554, 60, 90 (3 specimens) Brasil, Rio Grande do
Sul, Millstead; MCZ 22996-23000 (10 specimens) Brasil, Rio
Grande do Sul, Santa Maria, L. I. Price and T. E. White, 1936;
USNM 118180-1 (2 specimens) Brasil, Rio Grande do Sul, Santa
Maria, L. I. Price and T. E. White, 1936; USNM 121328 (1
specimen) Brasil, Rio Grande do Sul, Santa Maria, Inst.
Butantan.

LYSAPSUS LIMELLUS COPE

Savage and Carvalho (1953: 193) erroneously state that the

original description was based on Uruguayan material, when in

reality Cope (1862a: 156) says: "Habitat-Paraguay. Taken on

—.river " (leaving blank the name of the river) . Though

it was first described for Paraguay, Cope later himself (1862b:

126 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

351) cites it for Corumba, Steindachner I 1864: 262) for Caicara.
and Miranda Ribeiro. (1926:27) for Caeeres, all three localities
in Mato Grosso (Brasil). The latter author reports it also from
Rio Grande do Sul ( Brasil) , Uruguay, and Buenos Aires Province
(Argentina). Previously, Boulenger (1889; 246) had reported
the species from Resistencia, Chaco, Argentina and (1894: 347)
from Asuncion, Paraguay; Peracca (1895: 24) added Rio Apa,
Upper Paraguay, and Berg (1896: 161) Corrientes, Argentina.
Midler and Hellmich (1936: 25) and Vellard (1948: 169) had
specimens from the Province of Formosa ; Santa Fe can be added
to the Argentine localities (specimens in the Museo Argentino de
Ciencias Naturales). Outside of this general area of the basin of
the Paraguay, Parana, Plata and south of Brasil, Boulenger
(1898: 4) lists it for Beni, Bolivia; Baumann (1912: 122, 124,
161) for the Upper Amazon; Midler and Hellmich (1936: 26)
for the Marajo Island; Parker (1939:87) for Grao Para; Savage
and Carvalho (1953: 195) for the Para and Amazonas states.
Moreover, Parker (1935: 510) describes a closely related species,
Pseudis laevis, based on nine specimens from British Guiana and
one specimen from the Beni region (Bolivia) ; Savage and
Carvalho ( 1953 : 196) place this species in the genus Lysapsus.

In the Museum of Comparative Zoology and in the Chicago
Natural History Museum I have studied material from Paraguay
and Bolivia, in which some differences are noticeable between
specimens from the two localities. These differences permit the
species Lysapsus Umellus to be divided into two subspecies: L. I.
limellus Cope, from the Paraguay and Parana basin, and L. 1.
bolivianus sufsp. now from the Beni Province (Bolivia). 1 have
not seen material from Uruguay nor from southern Brasil ; any
such specimens belong to the subspecies L. I. limellus, but 1 pre-
fer not to assert this without having seen them. On the other
hand, Lysapsus hit vis can be considered as a third subspecies of
L. limellus, except for the Beni specimen, which undoubtedly be-
longs to L. I. bolivianus. Finally, it is possible that the specimens
from Para and Amazonas states belong to L. I. bolivianus; but
specimens from Rio Braneo (Brasil) that I saw in the United
States National Museum belong to L. I. laevis.

Lysapsus limellus limellus Cope

1862a Lysapsus limellum Cope, p. 155 ; 1862b, p. 351.

1882 Pseudis limellum Boulenger, p. 187: 1889, p. 246; 1894,

p. 347: Boettger, 1885, p. 242; Peracca, 1895, p. 24;

Berg, 1896, p. 161; Midler and Hellmich, 1936, p. 25

GALLARDO : SPECIES OF PSEUDIDAE 127

(part), fig. 8 ; Vellard, 1948, p. 169.

1953 Lysapsus limellus Savage and Carvalho, p. 194 (part).

Description. Adult female, MCZ 25834, Rio Pilcomayo, 15 miles
W. of Rio Paraguay, Chaco, Paraguay. Snout rather sharp,
nostrils 2 mm. from the tip of the snout. Back and ventral regions
with granulations. Cutaneous fringe of fingers insignificant. Di-
gital discs of the feet elliptic with rather sharp tips. Interdigital
membrane not very large; when the toes are adpressed it does
not form wide folds.

Coloration : A dark continuous line on the ventral internal
edge of the femur; posterior part of the femur without pattern.
Small black granulations on the ventral region of the femur.

Dimensions: Head and body 17 mm. Head length 6 mm. Head
width 6 mm. Head height 3 mm. Eye 2 mm. Snout 2 mm. Inter-
orbital space 2 mm. Elbow to third finger 10 mm. Femur length
12 mm. Tibia length 12 nun. Tibia width 2 mm. Heel to fourth toe

14 mm. Foot 10 mm.

Distribution : Paraguay and Parana Rivers and their affluents.
From Mato Grosso to Rio Tigre (Buenos Aires Province, Argen-
tina), as cited by Miranda Ribeiro (1926: 27). It has been re-
ported in Argentina from Formosa, Chaco, Santa Fe, Corrientes
and Buenos Aires. (The photo published by Cei, 1956 : pi. V, fig.
38, as of Pseudis mi nut us from Corrientes seems to belong to L. I.
limellus.)

Material studied: MCZ 25834 (1 specimen) Paraguay, Chaco,

15 miles W. of Rio Paraguay, Rio Pilcomayo, P. Willim, II-3 to
IH-3-1944; CNHM 42315-7 (3 specimens) Paraguay, Colonia
Nueva Italia, P. Willim. 1944; USNM 139277 (1 specimen) Para-
guay, San Bernardo, Lago Ypacaray (beach pools, C. J. D. Brown,
VIII-16-1956 ; USNM 139278 (1 specimen) Paraguay, Asuncion
Bay, Rio Paraguay nr. Asuncion, C. J. D. Brown, 1-2-1957 ;
USNM 132739 (1 specimen) Brasil, Mato Grosso, Paraguay River
above L. Gahiba-mirim, K. P. Schmidt, 1926; USNM 97146 (1
specimen) Brasil, Mato Grosso, Caceres; USNM 115975 (1 speci-
men) Brasil, Mato Grosso, Sao Luiz de Caceres, old cut-off of Rio
Paraguay, V-24-1909 ; USNM 132992 (1 specimen) Brasil, Mato
Grosso, Porto Esperanca.

Lysapsus limellus bolivianus subsp. nov.

1898 Pseudis limellum Boulenger, p. 4.
1935 Pseudis laevis Parker, p. 510 (part) .

128 BULLETIN: .MUSEUM OF COMPARATIVE ZOOLOGY

For diagnostic features see Table 2.

Description. Type, adult male, MCZ 17493, Reyes, Bolivia.
Snout shorter than in L. I. limcllus; nostrils very close to the tip
of the snout. Dorsal and ventral skin much granulated. Cutane-
ous fringe of fingers well devloped. Digital discs of the feet with-
out sharp tips. Interdigital membrane very well developed, forms
wide folds when toes adpressed.

Coloration : A continuous dark line on the internal edge of the
ventral aspect of the femur, as in L. 1. limellus, but also another
line more external, irregular and interrupted. Hind side of thigh
with a third dark line, interrupted, below the cloaca. Less abun-
dant small dark granulations on the ventral aspect of the femur.

Dimensions -. Head and body 18 mm. Head length 6 mm. Head
width 7 mm. Head height 4 mm. Eye 2 mm. Snout 1.5 mm. Tnter-
orbital space 2 mm. Elbow to third finger 9.5 mm. Femur
length 12 mm. Tibia length 12 mm. Tibia width 2 mm. Heel to the
fourth toe 15 mm. Foot 10 mm. Paratype, adult female. MCZ
10076, from the same locality, head and body 21 mm.

Distribution : The specimens studied come from the NW of
Bolivia, Beni Department, Amazonian drainage through the
Beni River and the Madeira. Thus it is probable that the speci-
men which Parker (1935: 510) ascribes to Pseuelis laevis, from
Santa Ana de Movrinas, Beni River, Bolivia, actually belongs to
L. I. bolivianus. (Boulenger, 1898: 4, reported L. limellus from
the same locality in Bolivia.) Also very likely to be this sub-
species are the specimens of Lower Madeira River in Amazonas
State and of Municipio of Monte Alegre in Para State, Brasil.
mentioned by Savage and Carvalho (1953: 195) and considered
by them to be L. limellus on account of the granulose skin (in
contrast to Pseuelis laevis which has smooth skin).

Material studied: MCZ 10076, 17493 (2 specimens) Bolivia,
Reyes, N. E. Pearson; MCZ 10077 (1 specimen) Bolivia, Lake
Rogagua, N, E. Pearson.

Lysapsus limellus laevis (Parker)

1935 Pseuelis laevis Parker, p. 510 (part).

Description. Adult male, USNM 137745, Fraseo, Boa Vista,
Rio Branco, Brasil. Snout short. Dorsal and ventral skin smooth.
Cutaneous fringe insignificant. Digital discs of the feet nearly
round. Interdigital membrane very well developed, forms wide
folds when toes adpressed.

<;allardo: species of pseudidae

129

Coloration : A continuous dark line on the internal edge of the
ventral aspect of the femur; more externally some dark spots in
line. A third dark line below the cloaca.

Dimensions : Head and body 21 mm.

Distribution : British Guiana, Rupununi Savannah and Paca-
raima foothills; in Brasil, Rio Branco near Boa Vista (this local-
ity is very near the British Guiana boundary) .

Material studied: USNM 137744-6 (3 specimens) Brasil, Rio
Branco, Boa Vista, Fraseo, J. D. Haseman, XI 1-5-1912.

L. I. limellus and L. I. laevis differ in the type of skin, the
shape of the snout, the extent of the interdigital membrane and
the coloration of the thigh ; the two are related by way of L. I.
bolivianus which has granular skin like L. I. limellus but in snout
shape, extent of interdigital membrane and in coloration is near
L. I. laevis, which differs from both by the extent of fringing on
the fingers.

L. I. laevis is restricted to the British Guiana and Rio Branco,
L. I. bolivianus to the Beni drainage and probably also the Ama-
zon, L. I. limellus to the system of the Paraguay. Parana and
Plata rivers.

Table 2. Diagnostic features of the subspecies
of Lysapsus limellus

Snout

Nostril from
tip of snout

Dorsal and
ventral skin

Cutaneous fringe
of fingers

Discs of
feet

Interdigital
membrane

Black lines on
ventral thigh

Posterior side
of femur

limellus bolivianus

Bather sharp Obtuse

Not very near Very near

Granulated

Insignificant

Elliptic with
rather sharp tips

Not very large,
not forming wide
folds

Internal edge with
a continuous line

Without design

Granulated

laevis
Obtuse
Very near

Smooth

Well developed insignificant

Elliptic without

sharp tips

Very well

developed,

forming wide

folds

Internal edge with

a continuous line

and a

discontinuous line

externally

With a third line

Nearly round

Very well
developed,
forming wide
folds

Internal edge with
a. continuous line
and some external
spots in line

With a third line

130 BULLETIN.: MUSEUM OF COMPARATIVE ZOOLOGY

Two of the species of the genus Pseudis are difficult to allocate ;
these are P. minutus Giinther, described on a specimen collected
by Darwin from "South America" (later regarded as coming
from South Brasil), and P. meridionalis Miranda Ribeiro, de-
scribed from Rio Grande do Sul, Brasil.

Pseudis minutus in some characters approaches P. paradoxus ;
such features are the general shape of the head, the lack of ter-
minal discs and the developmnt of the interdigital membrane ;
however, it resembles L. mantidactylus in size (40 mm. according
to Nieden) and in the presence of two vocal sacs (so specified by
Giinther in the original description, but Boulenger reports only
one). If Boulenger was right about the single vocal sac, it is very
probable that P. minutus is one of the southern races of P. para-
doxus; we must remember that Cope (1885: 187) reported P.
paradoxus from Rio Grande do Sul. Doubt was cast on this by
Boulenger (1885b: 298) who said confidently that Cope had the
species confused with L. mantidactylus. I think, however, that
this last is not at all probable since L. mantidactylus is a species
described by Cope.

Pseudis meridionalis, on the basis of its size (20 mm.) and the
resemblances to L. limellus mentioned by Miranda Ribeiro, could
very well be the representative in southern Brasil of the latter
species ; however, according to Miranda Ribeiro, the digital discs
are absent. (It must be remarked that what has sometimes been
identified as P. minutus or as P. meridionalis in many collections is
in reality L. mantidactylus.) All this makes desirable a better
study of the Pseudidae from southern Brasil, based on adequate
material.

SUMMARY

1) On the basis of both external and internal characters (pec-
toral girdle, digital phalanges), Pseudis mantidactyla Cope is
placed in the genus Lysapsus.

2) The color pattern of the ventral aspect of the thigh is suf-
ficiently constant in the species and subspecies of Pseudis and
Lysapsus to be taken as one of the useful characteristics in their
classification.

3) The distribution of the subspecies seems to be correlated with
the drainage system of the large South American rivers.

4) Six subspecies are defined for Pseudis paradoxus, three of
tliem new, and three for Lysapsus limellus, one new.

(JALLARDO: SPECIES OF PSEUDIDAE 131

ACKNOWLEDGMENTS

1 must thank the Consejo Nacional de Investigaciones Cienti-
ficas y Tecnicas de la Republica Argentina for the fellowship
given me for investigations on neotropical amphibians and for
study of the collections of the North American museums. I am
grateful also to Dr. A. S. Romer, Director of the Museum of
Comparative Zoology at Harvard University and to Dr. E. E.
Williams, Curator of Reptiles and Amphibians at this museum,
for the facilities afforded me during 1959-60; to the Council of
the American Museum of Natural History and the Karl P.
Schmidt Fund of the Chicago Natural History Museum for sup-
port given me and to Mr. C. M. Bogert, Dr. R. Inger, Dr. D.
Cochran, Dr. M. G. Netting, and Mr. N. Richmond for courtesies
and assistance offered me during my work in their several
museums; and to Dr. C. Gans of Buffalo University for supply-
ing data on material collected bv him in Bolivia.

LITERATURE CITED

Baumann, F.

1912. Brasilianische Batrachier des Berner naturhistorischen Museums.
Zool. Jahrb., 23 (2) : 87-172, fig. A., pis. 4-6.
Berg, C.

1896. Batracios argentinos, enumeracion sistematica, sinonimica y
bibliografica de los batracios. An. Mus. Nae. Buenos Aires, 5:
147-226.

BOETTGER, O.

188.1. Liste von Reptilien und Batrachiern avis Paraguay. Zeitschr.

Xaturwiss., 58: 213-248.
1892. Katalog der Batrachier-Sammlung ini Museum der seneken-

bergischen naturforschenden Gesellschaft. Frankfurt a. M.,

pp. 1-73.

BoULENGER, G. A.

1882. Catalogue of the Batrachia Salientia s. Eeaudata in the collec-
tion of the British Museum, London. Pp. i-xvi, 1-503, pis. I-XXX.

1883. Notes on little-known species of frogs. Ann. Mag. Nat. Hist.,
(5) 11: 16-19.

]88."ia. A list of reptiles and batrachians from the Province Rio Grande

do Sul, Brazil, sent to the Natural History Museum by Br. II.

von Ihering. Ann. Mag. Nat. Hist., (5) 15: 191-196.
1885b. Remarks on a paper by Prof. E. D. Cope on the reptiles of the

Province Rio Grande do Sul, Brazil. Ann. Mag. Nat. Hist., (5)

16: 294-298.

132 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1886. A synopsis of reptiles and batrachians of the Province Rio

Grande do Sul. Ann. Mag. Nat. Hist., (5) 18: 423-445.
1889. On a collection of batrachians made by Prof. Charles Spegaz-

zini at Colonia Eesistencia. Ann. Mus. Civ. Storia Nat. Genova,

(2) 7 (27) : 246-249, pi. II.
1894. List of reptiles and batrachians collected by Dr. Bohls near

Asuncion, Paraguay. Ann. Mag. Nat. Hist., (6) 13: 342-348.
1898. A list of the reptiles and batrachians collected by the late Prof.

L. Balzan in Bolivia. Ann. Mus. Civ. Storia Nat. Genova, (2)

19 (39): 1-6.
Cei, J. M.

1949. El ciclo sexual y el predomino de la espermatogenesis anual

continua en batracios chaquehos. Acta Zool. Lilloana, 8 :

527-544.

1956. Nueva lista sistematica de los batracios de Argentina y breves
notas sobre su biologia y ecologia. Invest. Zool. Chilenas, 3
(3-4) : 35-68, pis. I-IX.

Cochran, D. M.

1955. Frogs of southeastern Brazil. Bull. U. S. Nat. Mus., 206: i-xvi,
1-423, pis. 1-34.

Cope, E. D.

1862a. On some new and little known American Anura. Proc. Acad.

Nat. Sci. Philadelphia, 1862 : 151-159.
1862b. Catalogues of the reptiles obtained during the explorations of
the Parana, Paraguay, Yemiejo and Uruguay Rivers by Capt.
Thos. J. Page, USN ; and of those procured by Lieut. N.
Michler, U. S. Top. Eng., Commander of the Expedition con-
ducting the survey of the Atrato River. Proc. Acad. Nat. Sci.
Philadelphia, 1862 : 346-359.
1885. Twelfth contribution to the herpetology of tropical America.
Proc. Amer. Philos. Soc, Philadelphia, 22: 167-194, pi. 1.
Ditmars, Raymond L.

1942. The frogs of Paradox Pond. Animal Kingdom, 45: 51-55.
Fernandez, K. and M. Fernandez

1921. Sobre la biologia y reproduccion de algunos batracios argentinos.
I) Cystignathidae. Ann. Soc. Cient. Arg., 91: 97-140, pis. I-IIT.
Freiberg, M.

1942. Enumeration sistematica y distribution geografica de los ba-
tracios argentinos. Physis, 19: 219-240.
Gallardu, J. M.

1957. Las subespecies argentinas de B'tfo granulosus Spix. Rev. Mus.
Argentino Cien. Nat., 3 (6) : 337-374, pis. I-Y.

1958. Observaciones sobre el comportamiento de algunos anfibios
argentinos. Ciencia e Investigation, 14: 2H1-302.

Gans, C.

1956. Frogs and paradoxes. In Animaland (Staten Island Zool. Soc),
23 (4): 2-4, pi. 1.

GALLARDO : SPECIES OE PSEUDIDAE 133

1960. Notes on a herpetologicaJ collecting trip through the southeast-
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Garman, S.

1883. A species of Pseudis from the Rio Arassuahy, Brasil. Science
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GUNTHER, A.

1858. Catalogue of the Batrachia Salientia in the collection of the
British Museum, London: III-XVL 1-160, pis. I-XII.
Kenny, J. S.

1956. New locality records for the Paradox Frog. Jour. Trinidad Field
Nat. Club, 1956: 23.
Linnaeus, C.

1758, Systema naturae per regna tria naturae, secondum classes, or-
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Lutz, A.

1925. Batraciens du Bresil, I. Compt. Bend. Soc. Biol. Paris, 93 (21):
137-139.

1927. Notas sobre batraquios de Venezuela e da Ilha de Trinidad. Mem.
Inst. Oswaldo Cruz, 20 (1): 35-50, pis. 8-15.
Miranda Ribeiro, A. DE

1926. Notas para servirem ao estudo dos gymnobatraquios (Anura)
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MULLER L. AND W. HELLMICH

1936. Amphibien unci Reptilien I. Teil: Amphibia, Chelonia, Loricata.
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NlEDEN, F.

1923. Amphibia: Anura I. Das Tierreieh no. 46: 1-584.
Parker, II. W.

1933. A list of the frogs ami toads of Trinidad. Tropical Agriculture,
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1934. Some frogs and toads of Trinidad. Tropical Agriculture, 11 (5):
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1935. The frogs, lizards and snakes of British Guiana. Proe. Zool. Soc.
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1939. Resultats scientifiques des croisieres du navire-ecole beige "Mer-

cator" Vol. II, pt. V. Reptiles et amphibiens. Mem. Mus. Roy.

Hist. Nat. Belgique, (2) 15: 85-90.
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1895. Viaggio del dott. Alfredo Borelli nella Rep. Argentina e nell

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Savage, J. M. and A. L. Carvalho

1953. The family position of neotropical frogs currently referred to

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134 BULLETIN' : MUSEUM OF COMPARATIVE ZOOLOGY

SHREVE, B.

1959. A new Phyllomedusa from Bolivia (Salientia, Hylidae).
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Steindachner, F.

1864. Batra«hologische Mitteilungen. Verh. zool.-bot. Ges. Wien, 14:

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1867. Amphibien. Reise der osterreichischen Fregatte Novara urn die
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1942. Eelatorio da sexta excusao do instituto Oswaldo Cruz realizada
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1941. Mem. Inst. Oswaldo Cruz, 31 (3), Amphibia: 282-284.
Yellard, J.

1948. Batracios del Chac-o argentino. Acta Zool. Lilloana, 5: 137-174.
Wagler, J. G.

1830. Xatiirliehes System der Amphibien, mit vorangehender Classifi-
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den Zoologie. Miinehen, Stuttgart and Tubingen, 354 pp., 9 pis.

PLATE

Plate. Top: tadpoles of Lysapsus mantidactylus (above) and Pseudis
paradoxus (below). Bottom: adults of Pseudis paradoxus (left), Lysapsus
limeUits (center), Lysapsus mantidactylus (right).

Bulletin of the Museum of Comparative Zoology

AT HARVAED COLLEGE

Volume 125, No. 5

CORRELATIONS BETWEEN ECOLOGY AND

MORPHOLOGY IN ANOLINE LIZARDS

FROM HAVANA, CUBA AND SOUTHERN FLORIDA

By Bruce B. Collette

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

August, 1961

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MUSEUM OF COMPARATIVE ZOOLOGY
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Bulletin of the Museum cf Comparative Zoology

AT HARVARD COLLEGE

Volume 125, No. 5

CORRELATIONS BETWEEN ECOLOGY AND

MORPHOLOGY IN ANOLINE LIZARDS

FROM HAVANA, CUBA AND SOUTHERN FLORIDA

Bv Bruce B. Collette

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

August, 1961

No. 5 — Correlations Between Ecology and Morphology in
Anoline Lizards from Havana, Cuba and Southern Florida

By Bruce B. Collette1

INTRODUCTION

In many areas of the vast range of the iguanid genus Anolis
several species occur sympatrieally. There is, therefore, very
good opportunity to analyze the ecological separation of sympa-
tric species and to test the correlation of morphological with
ecological differences. Almost no use has been made of this op-
portunity up to the present. Oliver (1948) has indeed discussed
the ecology and morphology of Anolis on Bimini Ids. in the
Bahamas but he did not expressly correlate the two. Lundelius
(1957) appears to be the only worker who has carefully corre-
lated ecological differences with morphology (in two fence lizards
of the genus Sceloporus) .

In the present paper an attempt is made to correlate mor-
phology and ecology in five species of Anolis from Havana. Cuba.
The possible effects of introduced Anolis sagrei on Anolis caro-
livrnsis in southern Florida are also assessed.

MATERIALS

Over the past seven years nine trips of one to two weeks'
duration have been made to Havana, Cuba, to study the herpeto-
fauna. Two of the trips were in early September and the remain-
der were in the period from December 19 to January 4.

Field observations have been made on more than twenty
specimens of each of the following species: Anolis angusticeps
Hallowell, alutaceus (Cope), porcatus (Gray), and sagrei
( Dumeril and Bibron ) . The primary reason for using observa-
tions from a limited area was to eliminate effects of geographical
variation. In addition, one specimen each of A. argillaceous Cope
and A. equcstris Merrem, as well as three specimens of an appa-
rently new species of anole were taken. All observations on Cuban
species (except equestris) are based upon specimens personally
collected.

Additional specimens of A. equestris from other parts of the
province of Havana were used to supplement the single specimen
from the study area.

iRoom 71, United States National Museum. Formerly Department of Conserva-
tion, Cornell University.

138

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

THE STUDY AREA

The study area extends for about 1700 meters along the west
shore of the Rio Almendares in the city of Marianao across the
river from Havana. It is a low-lying strip no more than 200
meters wide. An abandoned road, the Carretera del Rio Almen-
dares, runs most of the length of the area and crosses the river
just below the Jardines de la Tropical. There is a fairly large
field in the northern part of the area and some other smaller
clearings in the woodland but most of the area is densely wooded.
A cliff about 150 feet high separates most of the area from
Marianao. There are some old ruins toward the southern end of
the area completely shaded over by large trees. Figure 1 shows
a stylized transect of the study area.

adult
porcatus

young porcatus^.

sagrei

u 1

Fig. 1. Stylized cross-section through the study area showing the niche
of five species of Anolis from Havana, Cuba.

THE SPECIES OF ANOLIS STUDIED

The ecology of each species of Anolis is described in this sec-
tion from personal field observations in the study area with sup-
plemental notes from the literature. Data are included on : where
each species has been collected; background color; body color;
method of locomotion ; population density ; and interspecies
contacts. .

Much of this information is summarized in Table I where
the species are listed in order of increasing " arbor eality."
As here used, the more time an animal spends in a tree or the
higher up in a tree it is, the more arboreal the animal is consid-
ered to be. This is an arbitrary method of distinguishing between
habitats without recourse to the creation of a new complex
terminology.

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 139

Anolis alutaceus (Cope): This small, slender, long-tailed anole
is brown with a mid-dorsal tan stripe and a white dewlap. It is
limited to a terrestrial niche in woodlands where it is found in
three types of habitat : in holes and at the base of the walls of
ruins ; in roekpiles ; and less often among the sticks and leaves of
the forest floor. Almost half the specimens were taken on the
ground and most of the others were in holes in the walls of the
ruins. Three were seen or taken in trees. When approached while
in holes, they may turn and retreat up to three feet into the hole
or they may hop from the hole down to the ground. Once they
land on the ground, they usually hop into the leaves and remain
there, often with the head and most of the body hidden beneath
the leaves. Part of the tail is usually visible if a careful search is
made. One specimen was seen on the low branches of a tree and
when chased it did not rely on its protective form and coloration
as others have. It used its long hind legs to hop from branch to
branch but did not make any attempt to run up, down, or along
the branches. None have ever been seen attempting to climb up or
down a wall.

An estimate of the relative abundance of alutaceus in the area
of the ruins was obtained on good collecting days. From 9 :40 to
11 :55 A.M. on December 29, 1955, seventeen specimens were seen
or captured. From 9 :40 to 11 :10 A.M. on December 23, 1958, six
alutaceus were collected. There were scattered individuals on the
ground elsewhere but they were more difficult to see and catch. No
fighting was seen between individuals of this species but this may
be due to the fact that they are not usually close together. In the
ruins, they may be only a few feet apart, but they are in separate
holes and cannot see each other. This species is limited to wood-
lands that are moderately damp and have a low light intensity.
The few body temperatures available indicate that it is active at
lower temperatures than the other species studied. Barbour
and Ramsden (1919: 154) also report that alutaceus is confined
to woodlands but note one unusual case of a specimen crossing a
hot, dusty road. Within the study area, alutaceus comes into
contact with angusticeps on the walls of the old ruins and with
porcatus, to a much lesser extent, near the base of some cliffs.

Anolis angusticeps Hallowell. A short-tailed, short-limbed,
long-headed anole which is gray or gray-brown with a pale
peach-colored dewlap. In the study area, it is limited to a wood-
land habitat and 17 of 23 specimens were collected on tree trunks
two to six feet from the ground. Three specimens were taken on

140 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the walls of the ruins and one was found on the ground. All were
taken in moderately moist areas of low light intensity. Individu-
als rest on tree trunks with the head pointed upwards and when
approached remain motionless for a short period but climb
quickly if the first attempt to collect fails. Although temperatures
were taken in the field on only four specimens, these show the
preference of this species for cooler areas. Oliver (1948: 7)
observed A. angusticeps chickcharncyi only on the upper branches
of light gray-colored trees on Bimini, Bahamas. In the study
area in Cuba, the species is found on gray or gray -brown tree
trunks. Here the abundance of angusticeps varies from year to
year. In 1957, five specimens were collected in 70 minutes on
the morning of December 24 and five more in 30 minutes on
the morning of December 27. In 1958, only two specimens were
taken in two weeks of collecting. In the study area, angusticeps
comes into contact with alutaccus on the walls of the ruins and
with porcatus on tree trunks near the edge of the woodlands.

Anolis argillaceus Cope. Only one specimen of this small stocky
lizard was taken in the study area. It was collected on December
24, 1957 at 10 :35 A.M. among some vines in the crotch of a tree
on the border of a clearing. When the vines co\ering the lizard
were moved, the lizard tried to run back under cover rather than
up the tree trunk as porcatus would, or down to the ground as
sagrei would have done.

Barbour and Ramsden (1919: 149) report this species as con-
fined entirely to eastern Oriente. This record constitutes a range
extension to the west of almost 500 miles.

Anolis equestris Merrem. A large handsome lizard with the
upper surface of the head developed into a magnificent bony
casque, it is usually a brilliant green and both sexes possess a pink-
dewlap. The only specimen taken in the study area, a juvenile,
was located at a height of six feet in a small tree in a field. Its
habitat as noted by Barbour and Ramsden (1919: 133-135) and
verified by Kane (personal communication) consists of orchards,
palm groves, and trees along roads. None have been seen on the
ground, and the majority of specimens have been seen at heights
of more than 15 feet in trees. Both sources note fighting on the
smooth trunks of the Royal Palm. Light and temperature inten-
sity is greater in these habitats than in the woodland habitat of
alutaccus and angusticeps and somewhat less than in the habitats
of sagrei and porcatus. Kane (personal communication) collected
one equestris that had partly eaten another anole but the speci-
men is not available for identification. In laboratory cages, I have

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 141

observed mating and other behavior taking place most often on
branches rather than on the cage bottom. In the study area,
equestris can come into contact with porcatus and possibly av-
gusticeps.

Anolis porcatus (Gray). A long-headed medium-sized anole
which has a purplish-pink dewlap. It is found everywhere in the
study area except in open fields and deep woodlands. Young are
common on small bushes and in tall grass in areas of high light
intensity. Adults are found on fence posts and tree trunks. The
young are usually green and the adults are usually brown. Cur-
rently, it is the commonest lizard in the study area. However,
since it is a species preferring edge areas, its great abundance is
due in part to man who by breaking up woodland areas and
planting trees in open areas, has created an ideal habitat for
porcatus.

In the study area, porcatus seems to be most active in morning
and afternoon with a period of decreased activity at midday. This
has been noted by Oliver (1955 : 133) for the related carolinensis
during the month of April in Florida where it is most active from
8-11 A.M. and again from 4-6 or 7 P.M. A. porcatus probably
spends the night in crevices or exposed on bushes or trees. Oliver
(1955: 134) has noted this behavior for carolinensis. Collections
made early in the morning revealed a high percentage of indi-
viduals under bark and in crevices in trees. Due to political con-
ditions in Cuba, no collecting could be done at night but one
specimen was observed asleep in the fold of a garden plant at
night.

Although it is difficult to estimate the abundance of porcatus
in the study area, the species seems to be found everywhere in
favorable habitats. Some indication of its abundance may be
gathered from the following field observations. On the trunk of
a large fig tree six specimens were collected, and the population
on the trunk from the ground to 15 feet up was estimated to be
at least 20 lizards. After the first few are collected from one tree,
the rest are disturbed, and it is almost impossible to catch all of
them. They frequently congregate in large numbers in limited
areas, a phenomenon not noted for the other anoles of the study
area. For example, under a sign with an area of six square feet,
at a height of seven feet above the ground, six to eight specimens
were noted at 1:30 P.M. Numbers of porcatus may be collected
in curled up fronds of broken palm branches that have lodged
in another tree. At least 20-30 juvenile to adult porcatus along

142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

with one Hyla septentrionalis and one Sphaerodactylus cinereus
were taken from one such palm frond. There were 15-20 in another
curled up palm frond. When the palm frond was unfolded, the
lizards immediately scurried off in every direction.

While territoriality has been noted for carolinensis in Rimini
(Oliver, 1955 : 111), in the study area, however, large populations
found on single tree trunks in December seem to indicate that
if territoriality occurs, it is not expressed to any considerable
extent at this time of year.

During this study, only one anole was observed feeding. A
small juvenile porcatus was seen sitting on a weed stalk about
four feet from the ground in a clearing. It crept slowly toward the
end of the stalk and grabbed a resting dragonfly.

In the study area, porcatus comes into contact with sagrei
frequently on fence posts. It also meets angusticcps in areas
bordering dense woodland and probably also contacts equestris
farther up in trees.

Since specimens of A. carolinensis Voigt in the Cornell Uni-
versity Herpetology Collection from the Fort Myers-Bonito
Springs area of southwestern Florida and from Key West were
examined in order to study the effects of populations of sagrei
upon carolinensis. a few remarks on this species are inserted here
to provide a proper comparison with A. porcatus.

Anolis carolinensis Voigt. This species is native to the United
States, is a smaller representative of the Cuban porcatus, and is
probably correctly considered as conspecific with porcatus
(Oliver, 1948: 12). I have observed this species only casually in
North Carolina and in Miami, Florida. From these observations
and from the literature, it is apparent that carolinensis occupies
both terrestrial and arboreal habitats though perhaps preferring
the latter. Duellman and Schwartz (1958 : 279) give the optimum
habitat in southern Florida as mesophytic hammocks, or in culti-
vated areas in gardens and shrubheiy around houses. The dewlap
is rose-pink to red and the body color varies within a wide range
of browns and greens. Over most of its range, carolinensis does
not come into contact with any other member of the genus.
However, on Key West and around several cities in southern
Florida, it does come into contact with introduced populations of
A. sagrt i.

Anolis sagrei (Dumeril and Bibron). This long-legged, short-
headed, alert anole has an orange-red dewlap and can change
color within a wide range of browns from pale tan to almost

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 143

black. It is the common lizard of the roadside and open field. The
juveniles and females are found mostly on the ground or a short
distance from the ground among weeds. Adult males are usually
found singly on observation perches at the top of woodpiles or
on fence posts. Schwartz and Ogren (1956: 98), Barbour and
Ramsden (1919: 143), Barbour (1904: 58, and 1914: 286), and
Oliver (1948 : 25) all report on its great abundance on the ground,
brush, fallen logs, and boards, fence posts, and piles of debris. In
the study area sagrei is second only to porcatus in abundance.
Evans (1938) notes that sagrei males defend territories from
other males of the same species ; no attempt was made to verify
this in the study area. However, two males were never taken on
the same fence post and only rarely on the same telephone pole.
When an attempt is made to approach sagrei on fence posts, they
quickly retreat to the ground and swiftly run away. This has also
been noted in the Cayman Islands by Grant (1940: 29) and on
Bimini, Bahamas, by Oliver (1948: 25). The males always face
downwards while on fence posts which indicates that their food
is terrestrial. Evans (1938: 103) notes that they make frequent
forays for insects in all directions from the observation perches
on fence posts. My field observations show that sagrei becomes
active somewhat later in the day than porcatus and has a peak of
activity in the middle of the day. Collections from 8 to 10 A.M.
have revealed that some individuals are partially active under
rocks. They probably spend the night under rocks and other such
cover. In the study area, sagrei comes into contact only with
porcatus at the base of trees and fence posts.

Anolis sp. Three specimens of a medium-sized white-throated
anole were also taken from the study area. They seem to resemble
A. Caroline nsis most closely. Nothing is known of their ecology
since they were not individually tagged in the field. They will be
treated in a subsequent paper.

Associated Herpetofauna. Since they are part of the total
ecology of the study area, the following list of herpetofauna is
presented. Amphibians: Rana catcsbeiana (one adult and many
tadpoles of this introduced species taken in backwaters of the Rio
Almenclares) ; Hyla septentrionalis (common in holes in trees) ;
EleutherodactyJus atkinsi atkinsi, E. euneatus, E. ricordii plani-
rostris1 (abundant under rocks throughout the study area).
Turtles: Pseudemys decussata (common on rocks in the Rio
Almendares). Lizards: Hemidactylus mabouia (at night near

irriiese Eleutherodactylus were determined by Mr. Benjamin Shreve.

144 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

house lights) ; Gonatodes fuscus (on logs and trees) ; Sphaero-
dactylus notatus (under debris throughout the study area) ;
Sphaerodactylus cinereus (under bark of trees throughout the
study area) ; Leiocephalus cubensis (common in edificarian situ-
ations near the study area) ; Amphisbaena cubana (a few indi-
viduals under debris in open areas). Snakes: Arhyton vittatum
(a few specimens under rocks) ; Tretanorhinus variabilis (a
juvenile under a car tire on the river shore) ; Alsophis angulifer
and Dromicus andreae (a few in a field several years ago before
it became overgrown) ; Typhlops lumbricalis (fairly common
under rocks) ; Tropidophis maculatus (few) and Tropidophis
pardalis (common) (both species under rocks and other debris).
Most of these species probably have little effect on the Anolis
in the study area. Tropidophis is the most important member of
the associated herpetofauna since both T. maculatus and T. par-
dalis include Anolis in their diets. Both A. alutaceus and A.
angusticeps have been found in the stomach of a T. maculatus.
In the laboratory, both species of Tropidophis feed on A. porcatus
and A. sagrei without hesitation. Captive Alsophis also feed on
Anolis but these snakes are not present in large enough numbers
to have much effect on the Anolis population.

SPECIES INTERACTION

Indications of fluctuations in populations of porcatus were
noted within the study area over a period of a few years. The
Carretera del Rio Almendares is lined, for part of its length, with
fence posts. In the years when all the grass around the posts was
cut down, sagrei was the most common lizard on the fence posts.
In years when there was tall grass in the field behind the posts,
and no grass along the road in front of the fence posts, some
sagrei and some porcatus were present on the posts. In the last
few years, some of the fence posts have taken root and furnish
shade in the areas. With this change, sagrei has decreased and
porcatus has increased in numbers on these living fence posts. In
nearby areas, tall grass sometimes completely surrounds fence
posts, and neither sagrei nor porcatus are present.

Although sagrei and porcatus are found on fence posts, they
usually occupy different positions. Typically adult male sagrei
are located facing downward on the lower part of the fence post
no more than a few feet from the ground. On the same fence post,
porcatus is found facing upwards on the upper part of the fence
post. If one species is absent, the other will tend to occupy the
entire fence post, but in all cases, sagrei faces downward and

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 145

porcatus upward. In the Cayman Islands this has also been noted
(Grant, 1940: 28) between A. sagrci ordinatus and A. maynardi
(considered conspecifie with porcatus by Barbour, 1937 : 119). It
is difficult to force porcatus to go down or sagrci to go up a fence
post. When a line of fence posts was approached, each sagrei ran
down the fence post into the leaves. A juvenile porcatus was
observed to run to the top of a fence post and then to jump to
two more fence posts in succession rather than move down the
fence post. If placed on the ground, porcatus seems uncomfortable
and it quickly moves to a nearby tree or fence post and climbs to
safety. "When approached by an observer, an adult porcatus
which is on a tree will warily circle upwards around the tree
putting the trunk between itself and the collector. This has also
been noted by Oliver (1948: 16) for the related A. carolinensis
lerneri on Bimini.

The Florida situation requires special attention. Populations of
sagrei now exist on Key West and adjacent keys and in several
mainland areas. The populations on the keys (A. sagrei stejne-
geri) have apparently reached there by natural means. The main-
land populations are recent introductions by man. Oliver (1950:
56) reports that Cuban A. s. sagrei have been introduced into the
Tampa area. A. sagrei ordinatus have been introduced from the
Bahamas into the Lake Worth area (Oliver, 1950: 56) and
reported from Miami (Bell, 1953: 63). Duellman and Schwartz
(1958: 283) refer the Miami populations to A. sagrei stejnegeri.
Willis (1953: 74) reports A. s. sagrei from Coral Gables and
Miami. At any rate, there are populations of the aggressive,
terrestrial sagrei in contact with carolinensis in a number of
areas in southern Florida. From studies by Oliver (1950),
Neill (1951), Bell (1953), and Duellman and Schwartz (1958),
it is apparent that the ecology of sagrei in regions of sympatry
with carolinensis does not differ noticeably from sagrei in the
study area in Cuba.

LAMELLAE

One of the most obvious differences in ecology of the
Anolis species studied is relative arboreality (Table I). (See the
definition above.)

The digital expansions of Anolis like those of geckos are un-
doubtedly adaptations to the generally arboreal habits of the
genus. Differences in the extent of arboreal habit might conse-
quently be expected to imply differing degrees of perfection and

146

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

efficiency of these arboreal adaptations. Certainly the width of
the digital expansion appears to correlate roughly with the
degree of arboreality. However, this is a character difficult to
make objective (See Fig. 2).

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Fig. 2. Feet of six species of Anolis showing the lamellae on the third
toe of the left hind foot, (a) alutaoeus, (b) angusticeps, (c) sagrei, (d)
Caroline nsis , (e) porcatus, (f) equcstris. Not to scale.

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 147

Another conspicuous character of the digital expansion may,
however, be quantitatively recorded — the number of lamellae.
The correlations of this character have therefore been examined.

Counts were made of the lamellae of the third and fourth toes
of the right front foot and the third toe of the right hind foot, of
the Anolis from the studj- area, plus samples of A. carolinensis
from Florida. These toes were selected because they have large
numbers of lamellae and therefore seem to be important in bear-
ing the weight of the lizard and are likely to show variation
between species, and because counts can be made accurately
since there is usually a distinct break between the lamellae of the
toes and the scales of the foot. Counts were not made on the fourth
hind toe because the distinction between lamellae and foot scales
is less evident there. Enlarged lamellae-like scales on the foot were
not counted. In the few cases where the proximal lamellae on the
toes were divided, the divided lamellae were counted as one.
Frequency distributions of the lamellae on each of the toes are
presented in Tables II and III.

From the data thus obtained, three factors appear to be corre-
lated with lamellae number. There is a positive correlation
between lamellae number and body size( Table IV) . This relation-
ship is understandable because as the length of the lizard in-
creases, the volume and presumablj* the weight increases as the
cube while the area of the toes increases as the square. The in-
creased number of lamellae in larger species compensates for the
loss in toe area relative to weight. Hecht (1952: 118) has also
shown a positive correlation between lamellae number and body
size in species of Aristelligcr. A second factor is that of sexual
dimorphism. In the study area, except for A. cqucstris and A.
alutaceus, male anoles have significantly more lamellae than fe-
males (Table IV). But this too may be a function of size since
(except for the two species mentioned above) male anoles are
larger than females (Table IV).1 However, this rule does not
appear to apply to Key West carolinensis in which, despite the
larger size of males, there is no significant difference in lamellae
number between males and females. Unless there is an ecological
difference between males and females, this is difficult to explain.
Thirdly, there appears to be a correlation between number of
lamellae and arboreality as it has been defined above. The more
arboreal species have more lamellae (Table IV). However, here

!But see ? angtustioeps compared with $ aagrei below.

148 BIIX.LETIN : MUSEUM OF COMPARATIVE ZOOLOGY

also there is some ambiguity. In general the more arboreal anoles
are also larger.1

Increased numbers of lamellae could be ascribed wholely to the
larger body size of arboreal species. However, there are several
lines of evidence that suggest that there may be a real rather than
a spurious correlation between arboreality and number of
lamellae.

In A. porcatus, the young are found in a bush-grass habitat
while adults are limited to a fence post-tree habitat. Superficial
examination indicates that the toes become wider with age but
more data is needed. There is no evidence of changes after hatch-
ing in lamellae number. The possibility of the phenomenon of

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SNOUT-VENT LENGTH

Pig. 3. Total number of lamellae on the third and fourth toes of the right
front foot and the third toe of the right hind foot of Anolis porcatus from
Havana, Cuba. The dots represent males and the crosses females.

iWithin a lizard genus, the more arboreal member of a group of gympatrie
species tends to be the larger. Within the study area, porcatus is larger than
sagrei; angusticeps is larger than alut actus : and equestris, the most arboreal
of all, is the largest one. Studies in progress on the gecko Sphaerodactylus also
show the arboreal cinereus to be larger than the terrestrial notatus. Smith (1946 :
204, 222) has shown that the arboreal Sceloporus oliraccus is larger than the
terrestrial S. undulatus. The significance of this trend is difficult to see.

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 149

wider variability in juveniles than in adults was tested in two
ways. The total number of lamellae on the three toes counted was
plotted against snout-vent length (Fig. 3). Until sex was taken
into consideration, it appeared that the mean number of lamellae
did increase with size classes, as found by Hecht (1952: 117) in
Aristelliger. But this was due primarily to sampling bias. There
are more males in the largest size groups and more females in the
smallest size groups so the differences in mean numbers of lamel-
lae are mostly an expression of the number of each sex in a
given size group.

Specimens of A. porcatus for which both exact ecological data
and lamellae counts are available fall into two ecological
niches: (1) bushes and grass; (2) fence posts plus trees. For
both sexes there is an increase in body size from the first to the
second group (Table V). The mean number of lamellae in the
bush-grass habitat is significantly lower than that for the fence
post plus tree habitat. Thus, with a shift in habitat, there is a
change in the distributions of body size and lamellae number
which seems to be due to selection acting against individuals with
low lamellae numbers. Selection acts not when a certain size is
reached, but when the young make the habitat shift to fence posts
and trees.

If lamellae number is plotted against snout-vent length for
sagrei, a similar picture is obtained (Fig. 4) . The main difference
between the lamellae-body size relations in the two species is the
greater separation between males and females in sagrei. However,
only the male sagrei make a habitat shift, while in porcatus both
sexes shift. Therefore, part of the male-female difference in
sagrei may be due to the larger maximum size that the male
reaches but part of it may be correlated with the fact that adult
male sagrei are found on fence posts while the females and juven-
iles of both sexes are limited to life on the ground.

Comparison of female angusticeps with female sagrei also sup-
ports the correlation of arboreality with increased numbers of
lamellae. Female sagrei are larger (mean 34.5mm, max. 44.4mm)
than angusticeps (mean 32.5mm, max. 38.9mm) but angusticeps
females have a larger mean and maximum number of lamellae.
Female sagrei are almost completely terrestrial while female
angusticeps, like the males, are arboreal.

A fourth case bears on the fpiestion of climbing ability and
lamellae number. A. carolinensis of Florida is closely related to
the Cuban porcatus and since it has no anole competitor over

150

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

L±J 60
<

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•

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• • • X X •

X XX
X X
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X X

1 1

I 1 1

20 25

30 35 40 45

55 60

SNOUT-VENT LENGTH

Fig. i. Total number of lamellae on the third and fourth toes of the
right front foot and the third toe of the right hind foot of Anolis sagrei
from Havana., Cuba. The dots represent males and the crosses females.

most of its range it is able to occupy all suitable anole habitats,
both terrestrial and arboreal. Both in lamellae number and body
size it is intermediate between two Cuban species — the arboreal
porcatus and the terrestrial sagrei (Table IV) . As discussed above,
sagrei has become established in a number of areas where caro-
linensis is present. Theoretically the aggressive, terrestrial sagrei
should tend to drive the more generalized carolinensis out of the
terrestrial part of its habitat. There then should be strong selec-
tive forces favoring those carolinensis with higher lamellae num-
bers, since they can better utilize the arboreal part of the habitat.
Counts on a small series of carolinensis from Key West, an area
of long contact with sagrei, are indeed higher than counts from
an area where they are not in contact (Fort Myers-Bonito
Springs), as shown in Table IV. These differences are significant
at the 99 per cent level for males and females (t values of 2.85
and 4.67 respectively). This is not conclusive evidence that the
suggested theory is correct because there is the possibility that
the lamellae numbers are correlated with greater body size, geo-
graphic variation in the latter character being the controlling

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 151

factor. Evidence for or against the theory can only be obtained
from study of carolinensis over a period of years in an area where
sagrei has been recently introduced.

TAIL LENGTH

Tail length was measured to the nearest tenth of a millimeter
from the vent to the tip of the tail. All specimens were carefully
examined and individuals with broken or regenerated tails were
excluded. Tail length was then divided by snout-vent length. A.
alutaceus has a much longer tail than the other species studied
(Fig. 5). This seems to be correlated with its use as a balancing

t — r— i — i — i — i — i — i r

ALUTACEUS

SAGREI

CAROLINENSIS

ANGUSTICEPS

PORCATUS

EQUESTRIS

i*i

1.5 2.0 2.5

Fig. 5. Ratio of body length to tail length in six species of AnoJis. The
horizontal line represents the range, the vertical line the mean, the filled-in
rectangle two standard errors on each side of the mean, and the open
rectangle one standard deviation on each side of the mean.

organ as this species hops along the forest floor. A. angusticeps
has a much shorter tail than the other species, possibly correlated
with the slow deliberate movements of this species. The other
tree-trunk dweller, porcatus, has a longer tail than angusticeps,
possibly because as a juvenile it makes daring leaps from leaf to
leaf and uses its tail as a balancing organ. All the arboreal species
have the base of the tail somewhat thickened for use as a prop in
climbing. This adaptation is not found in the terrestrial sagrei
and alutaceus. The tail of the former is compressed while that of
the latter is rounded and slender, well adapted for holding up in
the air as a balancing organ while the lizard hops.

152 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

HIND LEG LENGTH

Hind leg length was measured to the nearest tenth of a milli-
meter from the anterior insertion of the thigh to the tip of the
toenail of the longest toes with the leg extended at right angles to
the body. Dividing hind leg length into snout-vent length pro-
vided the ratio on which comparisons were based. The larger
arboreal species have shorter legs (Fig. 6). They move about

— i — r — i — i — i — i — i — i — i — i — I

ALUTACEUS

SAGREI pip

CAROLINENSIS
ANGUSTICEPS
PORCATUS
EQUESTRIS

J±L

J I I I I I I 1 I 1 I

1.5 2.0

Fig. 6. Eatio of hind leg length to body length in six species of Anolis.

more slowly in the trees than the terrestrial species do on the
ground. The long-legged terrestrial sagrei runs swiftly about in
fields while the long-legged alutaceus employs its long hind legs
in hopping on the forest floor. Lundelius (1957 : 80) reported that
the terrestrial Sceloporus undulatus has longer legs than the
arboreal S. olivaceus.

COLOR

Another adaptation to be considered is the ability of Anolis
to change color. The two terrestrial species, sagrei and alutaceus,
are limited to color and pattern shifts within a range of browns.
The ability to turn green would be of no positive advantage
against their brown backgrounds and would be selected against.

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 153

The arboreal angusticeps varies within a range of grays, gray-
browns, and gray-greens, the colors which are present on the tree
trunks of its damp woodland habitat. A. porcatus and carolinen-
sis have the ability to change color within a wide range of greens
and browns. Young porcatus are found among the leaves of
shrubs and bushes Avhere they are usually green. Adults are
usually found on the trunks of large trees in diffuse daylight
where they are usually brown.

The range within which a species can change color thus appears
to be adaptive to their ecological niche (Table I). Iladley (1929 :
110) notes that porcatus is green at night, brown in diffuse day-
light, and green in direct sunlight. Color changes are primarily
associated with changes in light, temperature, humidity, and
emotional state, but in most cases they result in the lizard match-
ing its background in nature (Van Geldern 1921: 81-87). An
exception is found where bright green porcatus (and carolinensis
in Florida) stand out against the brown of telephone poles and
isolated trees in situations modified by man. But perhaps this is
a habitat only recently colonized so there has been little time for
selection to act and few predators to provide selection pressure.

As for the completely arboreal equestris, both Kane (personal
communication) and Barbour and Ramsden (1919: 134) indicate
that this species is normally green. Hadley (1929: 112) notes that
equestris is normally green in diffuse light, the most frequent
condition in its niche.

PERITONEAL PIGMENTATION

Differences in the distribution of black pigment in the peri-
toneal cavity are present in the species of Anolis studied. Black
pigment is almost completely lacking in alutaccus. All the rest
have at least a pigmented parietal peritoneum. In angusticeps,
additional pigment is sometimes present on the ventral mesentery
that suspends the liver. Both porcatus and carolinensis have addi-
tional pigment on the mesocolon and mesoduodenum. The latter
also has pigment on the ventral mesentery of the liver. The pig-
ment distribution is similar in equestris but does not extend into
the most anterior portion of the peritoneal cavity. There is still
more pigment in sagrei; it covers the entire large and small in-
testines and extends onto the testes of the male.

Oliver (1948: 28) has noted similar conditions in the Bimini
A. angusticeps chickcharneyi, carolinensis lerneri, and sagrei
ordmatus, but offers no suggestions as to the possible significance

154 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

of these differences. There would appear to be a rough correla-
tion of the amount of peritoneal pigmentation with the amount
of radiation to which the lizard is exposed. The species of the
deepest woodland, alutaceus, lacks pigment and the other wood-
land species, angusticeps, has only a slight amount. The three
species of more open areas, equestris, carolinensis, and porcatus,
are exposed to greater radiation intensities and have more pig-
mentation. A. sagrei prefers open fields and has the most pigmen-
tation. It is possible that this pigmentation serves as protection
against harmful solar radiation as Klauber (1939: 77) has pro-
posed for reptiles in the southwestern United States. Even though
Bodenheimer (1954) and Hunsaker and Johnson (1959) have
indicated that the skin of some reptiles is impervious to various
forms of radiation, no alternative to Klauber 's explanation has
yet been proposed.

ACKNOWLEDGEMENTS

This paper and some of the ideas behind it have been discussed
with a number of people over the past five years and the author
is indebted to all of these. Mr. and Mrs. Raymond H. Collette, the
writer's parents made the study possible by providing transporta-
tion to Havana for collecting. Ernest E. Williams and A. Stanley
Rand, Harvard University, have given many valuable suggestions
in the preparation of this paper. Rudolph J. Miller, from the De-
partment of Conservation, Cornell University, has kindly drawn
Figures 1 and 2, and Edward C. Raney, Frederick R. Gehlbach,
Rudolph J. Miller and William A. Lund have encouraged the
study in discussions and by reading the manuscript. John Kane
has provided field observations and specimens of Anolis equestris
and Key West A. carolinensis.

SUMMARY

This paper has attempted to correlate ecology with morphology
in six species of Anolis from southern Florida and Havana, Cuba.
It is felt that with proper ecological data, valid correlations can
be made that can lead to an appreciation of the significance of
characters often used in taxonoinic analysis. Also, light is shed
upon the structural adaptations that allow related sympatric
species to occupy the same geographical area without facing
deleterious competition. It has been shown that selection has
acted so that lizards will usually match the color of their natural

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES 155

background. Examples have been shown to support the idea that
peritoneal pigmentation is connected with exposure to radiation.
The value of long legs to terrestrial lizards has been shown. Short
relative tail length has been correlated with arboreality. The more
arboreal members of a group of sympatric species have been
shown to be larger and have more lamellae than terrestrial
species. Data have been presented to support the contention that
increased numbers of lamellae are an adaptation to increased
arboreality.

LITERATURE CITED

Barbour, Thomas

1904. Batraehia and Reptilia from the Bahamas. Bull. Mus. Comp.

Zool., 46(3) : 55-61.
lit 14. A contribution to the zoogeography of the West Indies, with

especial reference to amphibians and reptiles. Mem. Mus. Comp.

Zool., 44(2): 209-359.

1937. Third list of Antillean reptiles ami amphibians. Bull. Mus. Comp.
Zool., 82 (2): 77-166.

Barbour, Thomas and Charles T. Ramsdex

1919. The herpetology of Cuba. Mem. Mus. Comp. Zool., 47(2):
71-213.
Bell, L. Xeil

1953. Xotes on three subspecies of the lizard Anolis sagrei in southern
Florida. Copeia, 1953(1) : 63.

BODENIIEIMER, F. S.

1954. Problems of physiology and ecology of desert animals. In
Biology of Deserts, Inst. Biol. London, pp. 162-167.

Duellmax, William E. and Albert Schwartz

1958. Amphibians and reptiles of southern Florida. Bull. Florida
State Mus., 3(5): 181-324.

Evans, Llewellyn T.

1938. Cuban field studies on territoriality of the lizard, Anolis sagrei.
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Grant, Chapman

1940. The herpetology of the Cayman Islands. Bull. Inst. Jamaica,
science series, 2 : 1-56.
Hadlev, Charles E.

1929. Color changes in two Cuban lizards. Bull. Mus. Comp. Zool.,
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Hecht, Max K.

1952. Natural selection in the lizard genus Aristelligcr. Evolution,
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llUNSAKER, BON II AND CLIFFORD JOHNSON

1959. Internal pigmentation and ultraviolet transmission of the integu-
ment in amphibians and reptiles. Copeia, 1959(4): 311-315.

156 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Klauher, Laurence SI.

1939. Studies of reptile life in the arid southwest. Bull. Zool. Soc.
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Lttndelitjs, Ernest L., Jr.

1957. Skeletal adaptations in two species of Soeloporus. Evolution,
11(1): 65-83.
Xeill, Wilfred T.

1931. A bromeliad herpetofauna in Florida. Ecology, 32(1;: 140-143.
Oliver, James A. *

1948. The anoline lizards of Bimini, Bahamas. Am. Mus. Novitates,

no. 1383: 1-36.
1950. Anolis sagrei in Florida. Copeia, 1950(1): 55-56.

1955. The natural history of North American amphibians and reptiles.
Princeton, N.J. Van Nostrand Co., Inc., 359 pp.

Schwartz, Albert and Larry H. Ogren

1956. A collection of reptiles and amphibians from Cuba, with the
descriptions of two new forms. Herpetologiea, 12(2): 91-110.

Smith, Hobart SI.

1946. Handbook of lizards. Ithaca, N.Y., Comstock Publ. Co., Inc.,
557 pp.
Van Geldern, Charles E.

1921. Color changes and structure of the skin of Anolis oarolinensis.
Proc. California Acad. Sci., (4) 10(10) : 77-117.
Willis, Edward T., Jr.

1953. Anolis sagrei in the Miami area. Herpetologiea, 9(2): 74.

COLLETTE: ECOLOGY AND MORPHOLOGY IN ANOLES

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Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE

Vol. 125, No. 6

CRANIAL ANATOMY OF THE CYNODONT
REPTILE THRINAXODON LIORHINIJS

By Richard Estes

With Two Plates

CAMBKIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

August, 1961

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Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE

Vol. 125, No. (i

CRANIAL ANATOMY OF THE CYNODONT
REPTILE TURIN AXODON LIORHINUS

By Richard Estes

CAMBEIDGE, MASS., U.S.A.
PEINTED FOE THE MUSEUM

August. 19H1

No. 6 --Cranial anatomy of the cynodont reptile
Thrinaxodon liorhinus

By Richard Estes1

The material discussed here was collected by the 1947 Uni-
versity of California African Expedition, Southern Section, at
Harrismith, Orange Free State. The locality data is as follows :
from Harrismith, six miles on the Bezuitenhout Pass road, then
two miles east to a farm, and one mile north to old stone corrals
on the point of a hill overlooking the river. These skulls were
picked up as nodules, which occur on the west and north slopes
of this point. The collectors were Dr. and Mrs. Charles L. Camp.
The locality is in the early Triassic Lystrosaurus zone, and is
designated University of California Vertebrate Locality number
V-36115.

The specimens dealt with below include two juvenile skulls,
which bear University of California nos. 42877 and 42878, and
two adult skulls, U. C. nos. 40466 and 42865. The juvenile
skulls were prepared by the use of ten per cent acetic acid
and treatment with Glyptal as described by Brink (1957b).
They are both crushed dorsoventrally, and lack the postorbital
bars. Badly crushed and broken lower jaws are also associated
with each of these skulls. U. C. no. 40466 is a fully adult skull ;
its preservation is perfect and undistorted, and very little is
missing. It was manually prepared by Mr. Martin Caulkin,
whose careful work on this specimen is gratefully acknowledged.
The other adult specimen is somewhat smaller, and is also well
preserved and nearly complete, but is slightly distorted. It was
also prepared in acid as described above to expose the basicran-
ium and the inside of the cranial cavity.

THE JUVENILE SKULLS

Brink (1955) has described an association of an adult and
a juvenile skull of Thrinaxodon liorhinus found together in the
same nodule. The juvenile skull is incomplete behind the orbits.
Juveniles are uncommon in the fossil record and the complete-
ness of the University of California specimens warrants a de-
scription supplementary to that of Brink.

The description is based on both skulls, and unless otherwise
stated, the characters are preserved on both specimens.

i Department of Biology. Boston University, and Research Associate, Museum
of Comparative Zoology, Harvard University.

166 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Snout

The premaxillae are missing' in no. 42878 and arc incomplete
in no. 42877. but in the latter the usual four incisors are present
on each side. The septomaxilla is well developed, and has a
posterior process inserted into the maxillo-nasal suture, a dorsal
intrafenestral process, and a ventral palatine process. The
prominent septomaxillary foramen is relatively larger than
that of the adult.

The nasals resemble those of larger individuals in being nar-
row anteriorly and pitted with tiny foramina. Posteriorly they
expand and articulate with the frontals. prefrontals, and
lacrimals, though this region of the nasals is not as much ex-
panded as in the adults.

The maxilla has the usual shape and contacts, and is also
strongly pitted with foramina. The teeth will be discussed below.

Skull Roof and Temporal Region

The frontals and parietals are relatively broader than those
of the adults, and the parietals lack a sagittal crest. The
temporal crests remain discrete throughout their length ; be-
tween them the flattened cranial roof is narrowest just posterior
to the nearly circular parietal foramen. The frontal differs in
both specimens from the condition characteristic of adult
Thrinaxodon liorhinus as follows. Each frontal sends a median
process into the midline suture between the nasals, forming a
wedge or arrow-shaped fronto-nasal suture instead of the usual
transverse one. The prefrontal-frontal and fronto-nasal sutures
thus form almost a straight line. Brink (in lift., 1958) has in-
formed me that his specimen also shows this condition, but to
a much lesser degree; thus the sutures mentioned above meet
at a greater angle.

The prefrontals are small and just touch the postorbitals. The
latter are fragmentary, but indicate that the posterior extent
of the postorbital was at the fronto-parietal suture or a little
posterior to it. The postorbital bars are broken away. The
jugals and squamosals closely resemble those of the adults.

The right quadrate is present on no. 42877, but it is badly
crushed, and only a tiny dorsal spike of the quadratojugal re-
mains in its groove in the squamosal.

ESTES : TURIN AXODON LIOI," I II NTS

167

dso

Fig-. 1. Dorsal view of restored juvenile skull of Thrinaxodon liorhinus;
X about 2.5. Abbreviations: aam, anterior ampulla; avsc, anterior vertical
semicircular canal; bo, basioccipital ; bs, basisphenoid ; cc, ems communis;
coch, cochlea ; cr, cochlear recess ; dso, dermosupraoecipital ; ec, ectopterygoid ;
co, exoccipital; ep, epipterygoid ; /./, jugular foramen; fo, fenestra ovalis;
fr, fenestra rotunda; fro, frontal; ic, internal carotid foramen; iv, interp-
terygoid vacuity; ju, jugal; la, lacrimal; m, muscle scar; ml, median line;
mx, maxilla; na, nasal; op, opisthotic; pa, parietal ; pam, posterior ampulla;
pf, prefrontal; pi, palatine; pm, premaxilla; po, postorbital; pr, prootic;
ps, parasphenoid ; pt, pterygoid; pvsc, posterior vertical semicircular canal;
<1J, quadra to jugal; qu, quadrate; sm, septomaxilla ; sq, squamosal; sr, sac-
cular recess; st, stapes; /, parasphenoid teeth; vf, venous foramen; ro,
vomer.

168 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Occiput

The occiputs of both juvenile specimens are crushed, but the
tabulars and dermosupraoccipitals are visible. In 42878 the
right half of the dermosupraoccipital is missing along what is
probably a suture, and 42877 shows a distinct suture in this
region, indicating that the bone was paired. The posttemporal
foramen is relatively a little larger than in the adult.

Lower Jaws

The lower jaws are badly crushed and broken in both speci-
mens ; no tooth counts could be made and no attempt has been
made to figure or restore either jaws or teeth.

The angulars are relatively a little larger and more flattened
externally than in the adults. The angular flanges are broken
away in both specimens.

Teeth

The juvenile thrinaxodons have a greater number of teeth
than do the adults, though the form of the tooth is very similar.

CD «,

Brink's juvenile specimen (Brink, 1955) shows seven teeth on
the right side. No. 42877 has seven teeth on both sides, and pos-
sibly eight on the left side. No replacement is noticeable in the
incisors, though all are quite loose in their sockets. The canines
are relatively small. The first three postcanine teeth on the left
side of no. 42878 are unicuspid and falciform, unlike the tricuspid
adult teeth of comparable position. On the right side of no. 42878
the fourth tooth is being replaced ; the tip of the replacing tooth
is barely emerging from the socket, although on the left side it
seems to be completely erupted. The replacing tooth on the
right is bicuspid, unlike the comparable tooth in the adult, which
is tricuspid. The sixth tooth on the right side is large and has
slipped down out of its socket. On the left side six postcanine
teeth are present, none of which seem to show any sign of re-
placement. However, the canine protrudes only slightly from
the alveolus and is apparently just erupting.

Teeth in adult specimens are much larger than those of the
juveniles. Replacement teeth in the juvenile are very little larger
than those replaced, indicating that further replacements would
be needed to bring the teeth up to the adult size. Brink (1955,
pp. 75-76) came to the same conclusion on the basis of his small

ESTES : THRINAXODON LIORHINUS 169

skull, and Crompton (If).")"), p. 665) has shown that some higher
cynodonts also have multiple (i.e. uon -mammalian) tooth re-
placement.

Palate and Basicranium

The palate and basicranium are fairly well preserved on both
specimens, but are more complete in no. 42877. The secondary
palate is as well developed as in fully adult Thrinaxodon
liorhinus; the maxillae and palatines meet on a long- midline
suture. A posterior palatine foramen is present on the maxillo-
palatine suture. The ectopterygoid is not preserved, but a
matrix-tilled cavity on no. 42878 indicates that this bone was
very small. There is a large palatal roof component of the
vomer, dorsal to the choanae, hut the median process which
would have reached to the secondary palate is broken away.

In contrast to adult cynodonts, the pterygoids enclose a small
interpterygoid vacuity on each side of the cultriform process
of the parasphenoid. Dorsally the basipterygoid processes are
smooth, and articulate on correspondingly smooth vertical faces
of the pterygoids, just forward of the internal carotid foramina ;
ventrally, the median ridges of the pterygoids curve mediad,
do not meet the cultriform process, and have a digitate suture
with the basipterygoid processes. The ventral keels of the
anterior portions of the pterygoids arc not continued posteriorly
onto the rounded basipterygoid tubera. The pterygoids continue
posteriorly, but are broken before reaching the quadrates.

The basisphenoid and parasphenoid are fused, except at the
anterior end of the basisphenoid dorsally, where a slight separa-
tion is present in the region of the trabecular attachment to the
basisphenoid, as described by Parrington (1935b, p. 400). The
tip of the cultriform process of the parasphenoid is sutured
anteriorly to the pterygoids, and the process extends posteriorly
between the interpterygoid vacuities. At this point the cultriform
process bears a ventral keel, and in the region between the
prominent internal carotid foramina, it expands and bears five
small teeth on a roughened area. From this area, the wings of
the parasphenoid expand and pass back over the basioccipital in
a squamous suture, the full extent of which is obscured by
breakage. There is a small gap between the basioccipital and
basisphenoid, which was undoubtedly tilled with cartilage.

The basioccipital is a hexagonal bone, and bears paired oval
depressions on the ventral surface, which were interpreted by

170

IU'LLETIX: MUSEUM OF COMPARATIVE ZOOLOGY

Parrington (1946b, p. 186) as attachments for the rectus capitis
anticus muscles. These depressions are relatively much deeper
than those of larger specimens. In no. 42877, the left one has a
foramen opening' into its lateral wall, facing medially and a
little posteriorly. This foramen opens into a canal, which disap-

Fig. 2. Ventral view of restored juvenile skull of TJirina.rodon liorhinus;
X about 2.5; for abbreviations see Fig. 1.

pears into the unossified area between the prootic and basioccipi-
tal. The foramen and its canal are formed wholly within the
basioccipital. This is certainly the same structure as that de-
scribed by AVatson (1913, p. 220) in Diademodon, and is, as he
suwge.-ts, probably venous. On the posterior slope of each muscle
pit is a smaller foramen which is directed towards the occipital
condyles. These are also visible in the adult (see PI. ], tig. 2).

ESTKS : THRINAXODON LIORHINUS

171

In addition, a tiny foramen is present on the opisthotic about
midway between the posterior borders of the jugular foramen
and the fenestra oval is.

( )tic Region

The otic region is shown in detail in Figure 3B. Several addi-
tions may be made to Parrington's account of the fenestra oralis
region of Thrinaxodon. Of particular interest is the fact that he
describes the fenestra oralis as confluent with the "unossified
region" (to be discussed below), which for the juveniles in-
cludes an area separating basioccipital from basisphenoid and
prootic, except for a tiny contact with the latter at the anterior
border of the cochlear recess (see below) in both adult and

Fig. 3. A. Eestoration of left inner ear of juvenile Thrinaxodon liorhinus;
superointernal view, X about 7. Dotted line indicates position of horizontal
semicircular canal.

B. Restoration of left otic region of juvenile Thrinaxodon liorhinus;

itral view, X about 7. The top of the figure is anterior. Parasphenoid

vent

itted to show underlying structures. For abbreviations see Fig. 1.

juvenile. Figure 3B shows that in the juvenile, a process of the
opisthotic almost completely closes the fenestra ovalis anteriorly ;
thus about three-fourths of the fenestral border is formed by the
opisthotic, the remainder by prootic. A parasphenoid contribu-
tion to the fenestral border is small, if present, and the fenestra
dors not reach the basioccipital.

172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Posteriorly, the fenestra oralis is confluent with the jugular
foramen through a distinct channel, which lies entirely within
the opisthotic and emerges within the jugular foramen near its
external opening. It was apparently a complete canal in life.
and must represent the fenestra rotunda. The presence of a
fenestra rotunda was first demonstrated in therapsids by Simp-
son (1933, p. 289) in Nythosaurus. Olson (1944, p. 25) suggested
that the perilymphatic duct in therapsids opens into the jugular
foramen. These juvenile specimens demonstrate that this was
the case in Thrinaxodon liorhinus, and that a fenestra rotunda
was present in cynodonts more primitive than Nythosaurus.

Anterior to the fenestra ovalis, a deep recess is present in the
skull base. It is formed principally in the basioccipital, except
for its anterolateral and posterolateral corners, which consist of
prootic and opisthotic, respectively. Apparently a thin cartilage
coating was present on the interior of this recess., but its definite
shape and relatively smooth inner surface indicate that it
housed a soft structure and was not cartilage filled. The re-
cesses were probably covered by parasphenoid in life, as in the
adult, but breakage has now exposed them on no. 42877. The
recess is partially roofed laterally by a small process of the
prootic, which tends to separate it from the saccular recess (see
below, and Fig. 3B). The recess in the basioccipital is present
only as a slight concavity in the adult. In no. 42865 (a young
adult in which the brainease was prepared by acid) the area is
very slightly concave, but no distinct recess is present. The
conformation of the fenestra ovalis is very similar, in the young
adult, to that described by Parrington (1946). However, in no.
42865, the anterolateral notch, in the large opening described by
Parrington as the fenestra ovalis, has a smooth, finished edge
internally, and is confluent with the area immediately ventral
to the anterior ampulla. This particular notch is probably the
ventral edge of the saccular recess, while the recess in the
basioccipital is certainly a receptacle for a cochlear apparatus.
Relative reduction of the basioccipital recess in the adult is prob-
ably a result of early development of the ear region of vertebrates
and very little increase in size as adult size is reached.

The jugular foramina are large, formed between the exoccipi-
tal and opisthotic, and on their concave posteromedial borders a
pair of small foramina are present. These are confluent with
the condylar canal, and carried the hypoglossal nerves.

The stapes is roughened and unfinished proximally, and evi-
dently bore a heavy cartilage plug, fitting into the slightly
bevelled sides of the fenestra ovalis. It was perhaps similar to

ESTES: THRINAXODON LIORHINUS 173

the ossified plug1 described by Parrington (1955, p. 14) in
Scylacops capensis. Only about one-half of the articular end of
the stapes covers the fenestra ovalis. The remainder projects
into, and partly over, the opening- into the so-called "unossified
region" (Parrington, 1946b, p. 185) and thus comes near the
lateral wall of the cochlear recess. The distal end of the stapes
is unossified. There is a relatively large stapedial foramen, and
the posterior limb of the stapes is slenderer than the anterior
limb, the latter condition resembling that of the adult. The
fenestra ovalis is separated from the "unossified region" by an
anterior process of the opisthotic. Anteriorly, the vestibule of
the fenestra ovalis is confluent with the cochlear and saccular
recesses.

Restoration of the Inner Ear

A restoration of the inner car of the juvenile (Fig. 3A) shows
the position and relationships of the fenestra ovalis, cochlea, and
fenestra rotunda. It is very similar to that figured by Olson
(1944, fig. lOd) for the adult Thrinaxodon, differing chiefly in
the relatively thicker semicircular canals, exit for the fenestra
rotunda, and presence of the cochlea. So far as the bony canals
are concerned, the ear structure is comparable with that figured
by Simpson (1933, fig. 3) for late cynodonts. and later, for the
monotreme Tachj/glossus (Simpson. 1938. fig. 2).

Fnossified Area of the Skull Base

The unossified area in the skull base of therapsids varies in
extent in the various groups. Usually there is an unossified
region between basioccipital and basisphenoid, and this may
extend into the ear region. In Thrinaxodon liorhinus the con-
tact between prootic and basioecipital-basisphenoid is mostly
unossified in both adults and juveniles. There is almost a con-
tact anteriorly between the prootic and basisphenoid in the
juveniles (see Fig. 3B), but otherwise the area was cartilage
filled. In the young adult skull mentioned above (no. 42865),
the unossified gap between the basisphenoid and basioccipital
is very small. From the prootic-basioccipital contact posteriorly,
the side wall of the inner ear capsule of this specimen is unossi-
fied back as far as the fenestra ovalis. At this point the latter
is separated from the unossified area by the process of the
opisthotic mentioned above in the section on the otic region. This
process is also present, though less discrete, in the fully mature
skull discussed below.

174 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Juvenile Characters of the Small Specimens

The restorations given in Figures 1 and 2 show that the two
small specimens differ in few respects from adult Thrinaxodon
liorhinus. The differences which do occur are probably the re-
sult of juvenility. These differences are discussed below.

The ossification is light, more so than might be expected in the
adult of a smaller species. The bone is very cancellous and
translucent.

All sutures are clearly visible. This is also often true of the
adults, but the latter have well-knit, occasionally complex and
inter-digitated sutures, while those of the small specimens are
relatively more open and have less complication.

The presence of small interpterygoid vacuities can be ex-
plained by juvenility of the specimens. Presumably the pterygoid
musculature was not yet well developed ; in the adult, these mus-
cles obliterate the vacuities by appression of the medial flanges of
the pterygoids against the midline. In the adult specimen dis-
cussed below (see also PI. 1, fig. 1), the former position of the
interpterygoid vacuities shows as two slim grooves on each
side of the midline.

The presence of parasphenoidal teeth in a therapsid is unex-
pected. Palatal teeth on pterygoids and palatines are found in
gorgonopsians, as well as in other groups of mammal-like reptiles,
and Vaughn (1958) has described snb-sphenoidal teeth in a
small pelycosaur. These teeth may be more consistently present
in therapsids than previously supposed. Perhaps they have
passed unnoticed owing to removal by mechanical preparation,
or obscured in the older individuals by appression of the
pterygoids to the midline.

Differences in tooth number have been considered to be taxo-
nomically significant, but individual variation between right and
left sides in both juveniles and adults vitiates its utility. An
example of ontogenetic variation is provided by the comparison
of the greater number of juvenile than adult teeth in Thrinaxodon
liorhinus with the converse situation in Galesaurus planiceps
i\i\<xncy, 1938, p. 512).

In animals which, as adults, possess well-developed sagittal
crests, the lack of development of these crests in juveniles is a
frequent phenomenon, both in reptiles and mammals. Thus, the
weak temporal crests, relatively large parietal foramen, and
flattened skull table between the temporal crests of these small
specimens indicate their juvenility.

ESTES: THRINAXODON LIORHINUS 175

The principal difference between the University of California

specimens and Brink's juvenile one (Brink 1955, p. 73) is in
the degree of development of the anterior projections of the
frontals. These spines, which thrust a wedge anteriorly between
the nasals, are also found in Glochinodontoides, Platycraniellus,
and Galesaurus. Tins is apparently a variable character in the
juveniles of Thrinaxodon liorhinus, disappearing in the adults
owing to further growth of nasals and frontals.

The other differences between adult and juvenile, pointed out
by Brink, are present in the University of California specimens,
though the presence of incomplete postorbital bars cannot be
determined in this material. The relatively very large frontal
of the juvenile is separated from the orbital margin by a very
narrow meeting of the postorbital and postfrontal. The posterior
margin of the secondary palate is at the level of the fifth tooth in
the juvenile rather than the third as in the adult. The latter
may he explained by the greater relative growth of the snout in
the adult, and the greater number of anterior teeth in the
juveniles.

THE ADULT SKULL

The specimen discussed here also came from the Lystrosaurus
zone near Harrismith. and has been designated U. ('. no. 40466.
Plates 1 and 2 were prepared some years ago by Mr. Owen Poe,
Staff Illustrator for the Museum of Paleontology, University of
California. They are included here because no detailed figures
of actual specimens of T. liorhinus have ever been published.
The general anatomy of the adult Thrinaxodon skull has been
elucidated by Parrington (1946b), and Olson (1944) has dis-
cussed the ear region, but this specimen shows some additional
points of interest.

Foramina of the Snout

The maxilla bears a row of four to six large superior labial
foramina along its ventral exterior surface. Other more dorsally
placed foramina are more numerous, smaller, and open anteriorly,
though some of the posterior ones are directed backwards. The
anterior parts of the nasals also bear numerous similar foramina.
Watson (1931, p. 1200) has suggested that in Ericiolacerta
these foramina indicated either a mobile prolongation of the
snout or highly developed sense organs such as a rhinarium or

176 BULLETIN: MUSKUM OF COMPARATIVE ZOOLOGY

vibrissae. Brink (1957a, p. 86) extends this interpretation to
Diademodon, as does Attridge (1956, p. 67) for the therocephal-
ian Mirotenthes. The broadly ancestral position of theriodonts
to mammals may justify an interpretation of these foramina as
indication of a rhinarium-like structure. It should be pointed
out, however, that an equally vascular bone surface is present in
some lizards. Tupinambis (Teiidae) shows near identity with
Thrinaxodon in the shape, number, and distribution of these
foramina. In addition, Tupinambis shows a dendritic pattern of
vascular grooves emanating from many of the more dorsal maxil-
lary foramina. These arborizations resemble those described by
Brink (1957a) for Diademodon, though they trend dorsopos-
teriorly rather than anterodorsally as in Brink's specimen.
Tupinambis also has an enlarged superior labial foramen, which
opens forward on the maxillary surface near the posterior border
of the external naris and is frequently continued into the latter
by a shallow gutter. This condition duplicates that of Ericio-
lacerta, as described by Watson.

Van Valen (1960, p. 306) mentions this vascularization of the
snout region in Tupinambis, but suggests that the foramina
"from their positions and relative development to be mostly
related to the development of the teeth." On the contrary, these
external maxillary (superior labial and lateral ethmoidal)
foramina of lizards transmit only nerves and blood vessels
serving cutaneous structures. Dorsally, the foramina perforat-
ing the nasals and dorsal part of the nasal processes of the
maxillae transmit cutaneous branches of the lateral ethmoidal
nerve. These serve highly vascularized and richly innervated
skin thickenings which surround the cartilaginous nasal tube
(Oelrich. 1956, p. 88). All of the snout skin is firmly attached
to the underlying bone, with the exception of the above-mentioned
thickened area, which is slightly motile (ibid., p. 87), though
no more so than in any other lizard.

The labial foramina transmit cutaneous branches of the
superior alveolar nerve and maxillary artery, both serving the
skin of the lower snout and the lip (ibid., pp. 62-63). The latter
is very weakly developed in all lizards. Those branches of the
maxillary artery and superior alveolar nerve which serve the
teeth are transmitted through ventral ly and internally opening
foramina in the palatal shelf of the maxilla, and are completely
separate from branches serving cutaneous structures.

ESTES : THRINAXODON LIORHINUS 177

The similarity of bone vascularization in Thrinaxodon and
Tupinambis suggests, then, that these foramina in themselves
may not be sufficient evidence to indicate the presence of asso-
ciated extensive secretory, sensory, or muscular structures in
Thrinaxodon or indeed in any other theriodont. Presence of
numerous large foramina on the lower part of the maxilla near
the alveolar horder would, on the other hand, seem to preclude
the presence of an extensive movable muscular cheek and lip.
The vascular and nervous structures associated with these
foramina would probably serve skin fairly closely united with
the bone, for extensive movement of this skin would be disad-
vantageous. A muscular cheek, then, is probably to be corre-
lated with the development of a posteriorly placed, single, ex-
ternal, infraorbital foramen as seen in mammals. So far as I am
aware, no such structure, or even a tendency for such a grouping
or codification of nervous and vascular snout structures is pres-
ent in any theriodont.

in summary, this discussion does not suggest that rhinarium,
hair, muscular cheeks and mobile lips were not present at all in
some therapsids; rather, that at least some of the evidence which
has been adduced for the presence of these or similar structures
is necessarily inconclusive. An excellent summary of this evi-
dence is given in Van Valen (1960).

Otic Region

The stapes of this adult specimen (no. 40466) was removed
from the left side and cleaned, and the region of the fenestra
ovalis carefully prepared (PI. 2, figs. 2-5). The stapes resembles
in a general way that described by Parrington (1955, p. 14.
fig. 8) for Scylacops capensis. It consists of a round head, the
posterior two-thirds of which fits over the fenestra ovalis. This
head is concave, contrasting with the bevelled plug of Scylacops
and also with tin1 rounded head of the stapes of the juvenile
Thrinaxodon. The head is continued ventrolateral^ by two
slender arms enclosing a large stapedial foramen, which opens
dorsoventrally rather than anterolaterally as in Scylacops. The
posterior arm is more slender, as in the juvenile. On the distal
end of the bone a strong dorsal process curves posteriorly, so that
the upper portion is visible ventrally. There is a smooth articula-
tion with the quadrate rather than a pronounced boss as in
Scylacops.

178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Temporal Musculature

Of special interest in this specimen are thin, bony sheets,
present on both sides, and attached to the slightly thickened
ventral border of the epipterygoid. These are visible in Plate 1,
figures 1 and 2 as irregular sheets occluding the temporal vacui-
tics. These sheets trend posteroventrally and are wrinkled and
distorted. They are interpreted as ossified fasciae, and thus
directly indicate attachment of a broad, muscular, aponeurosis.
Ventrally on the right side, this ossified fascia approaches
the posterior bones of the lower jaw near the anterior part of
the angular. The broad posterior attachment of the fascia along
the anteroposterior ventral edge of the epipterygoid excludes
the possibility of the fascia reflecting the pterygoid musculature.
It agrees, however, with Parrington's placement (1955, p. 22,
fig. 11) of the temporal muscles, and probably reflects their
lower attachment. These muscles probably inserted on the pos-
terior bones of the lower jaw (perhaps in the fossa between the
surangular and prearticular ) as their homologue, the adductor
externus group, does in Sphenodon and other reptiles.

SUMMARY

Two small skulls of Thrinaxodon liorhinus, a cynodont reptile
from Lower Triassie Karroo bed- of South Africa, show differ-
on :es from large specimens which lead to interpretation of the
former as juvenile. They provide the first record of para-
sphenoidal teeth in a therapsid, and demonstrate for the first
time the presence of small interpterygoid vacuities in cynodonts.
The otic region shows a fenestra ovalis almost wholly surrounded
by prootic and opisthotic; a large recess in the basioccipital
anterior to the fenestra ovalis probably housed a cochlea. A
distinct channel connecting fenestra ovalis with jugular foramen
is undoubtedly a fenestra rotunda. In a fully adult specimen,
ossified fasciae give direct evidence of temporal muscle attach-
ment. The presence of a highly vascularized maxilla in
theriodonts is probably insufficient evidence in itself to indi-
cate an extensive sensory or muscular cheek and snout region
of mammalian type.

ACKNOWLEDGEMENTS

I wish to thank the following people for numerous helpful sug-
gestions and critical reading of the manuscript: C. L. Cam]).
Bryan Patterson, and J. T. Gregory. Special thanks must go

ESTES : TIIRIXAXODOX LIORHIXUS 179

to Dr. II. 15. S. Cooke for his advice and assistance in acetic acid
and Glyptal techniques. I am also grateful to Dr. A. S. Brink
for providing a sketch of the juvenile specimen described by him.

LITERATURE CITED

Attridge, John

1956. The morphology and relationships of a Complete therocephalian
skeleton from the Cistecephalus zone of South Africa. Proc. Roy.
Sue. Edinburgh, vol. 66, ser. B, pt. 1, no. 4, pp. 59-93, 9 figs.,
2 pis.
Brink, Adrian S.

1955. Note on a very tiny specimen of Thrinaxodon liorhinus.

Palaeontol. Afrieana, vol. 3. pp. 73-76, 1 fig.
1957a. Speculations on some advanced mammalian characters in the
higher mammal-like reptiles. Palaeontol. Afrieana, vol. 4, pp.
79-96, 5 figs.
1957b. On the usas of Glyptal in paleontology. Palaeontol. Afrieana,
vol. 4, pp. 124-130.
Crompton, A. W.

1955. On some Triassic cynodonts from Tanganyika. Proc. Zool. Soe.
London, vol. 125, pp. 617-669, 15 figs.

Of.lrich, Thomas M.

1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae).
Univ. Mich. Misc. Publ. Mus. Zool., no. 94, pp. 1-122, 59 figs.

Olson, Everett ( !.

1944. Origin of mammals based upon cranial morphology of the
therapsid suborders. Spec. Pap. Geol. Soc. Amer., no. 55, ix +136
pp., 27 figs.
Parrington, F. R.

1935. A note on the parasphenoid of the cynodont Thrinaxodon
liorhinus Seeley. Ann. Mag. Nat. Hist., ser. 10, vol. 16, pp.
399-401, 1 fig.
1946. On the cranial anatomy of cynodonts. Proc. Zool. Soc. London,

vol. 116, pp. 181-197, 10 figs.
1955. On the cranial anatomy of some gorgonopsids and the synapsid
middle ear. Proc. Zool. Soc. London, vol. 125, pp. 1-40, 15 figs.
Kigxey, Harold W.

1938. The morphology of the skull of a young Galesaurus planiceps
and related forms. Jour. Morphol., vol. 63, pp. 491-529, 8 figs.,
6 pis.
Simpson, George Gaylord

1933. The ear region and the foramina of the cynodont skull. Amer.

Jour. Sci., ser. 5, vol. 26, pp. 285-294, 5 figs.
1938. Osteography of the ear region in monotremes. Amer. Mas.
Xovit., no. 978, pp. 1-15, 7 figs.
Van Yalen, Leicii

1960. Therapsids as mammals. Evolution, vol. 14, no. 3, pp. 304-313.

180 BULLETIN* : MUSEUM OF COMPARATIVE ZOOLOGY

Vaughn, Peter P.

1958. A pelycosaur with subsphenoidal teeth from the Lower Permian
of Oklahoma. Jour. Washington Acad. Sci., vol. 48, pp. 44-47,
1 fig.
Watson, I>. M. S.

1913. Further notes on the skull, brain, and organs of special sense of
Diademodon. Ann. Mag. Nat. Hist., ser. 8, vol. 12, pp. 217-228,
5 figs.
1931. On the skeleton of a bauriamorph reptile. Proc. Zool. Soc.
London, 1931, pp. 1163-1205, 27 figs.

PLATES

Plate 1
Fig. 1. Thrinaxodon liorhinus, adult specimen, dorsal view of U. C. no.
40466; X 1.5. Figure by Owen J. Poe.
Fig. "2. The same, ventral view.

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Bulletin of the Museum of Comparative Zoology

AT HAEYAED COLLEGE

Vol. 125, No. 7

TWO SYMPATRIC CUBAN ANOLES OF THE
CAROLINENSIS GROUP

RODOLFO RUIBAL

Division of Life Sciences
University of California, Riverside

and

Ernest E. Williams

Museum of Comparative Zoology
Harvard University

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

August, 1961

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with THE

MUSEUM OF COMPARATIVE ZOOLOGY
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Bulletin of the Museum of Comparative Zoology

AT HAEVARD COLLEGE

Vol. 125, No. 7

TWO SYMPATRIC CUBAN ANOLES OF THE
CAROLINENSIS GROUP

RODOLFO RUIBAL

Division of Life Sciences

University of California, Riverside

and

Ernest E. Williams

Museum of Comparative Zoology
Harvard University

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

August, 1961

No. 7 — Two sympatric Cuban anoles of the carolinensis group

By
rodolfo ruibal

and

Eknest E. Williams

INTRODUCTION

The carolinensis group of the West Indian iguanid genus
Anolis consists of forms sufficiently closely related that most or
all of them have been considered subspecies of one species. The
described forms occur both on the mainland of North America
and on the Caribbean islands: carolinensis in the southeastern
portion of the United States; lerneri, orunneus, smaragdinus on
the Bahamas ; fairchildi on Cay Sal ; longiceps on Navassa ; may-
nardi on Little Cayman ; porcatus on Cuba ; and allisoni on the
Islas de la Bahia off the coast of Honduras and on Half Moon
Cay off British Honduras. Only one representative of the group
has ever been reported from any one locality. Thus it has always
been assumed that porcatus was the single Cuban representative
of the carolinensis group.

Some years ago an examination of the specimens under the
name Anolis porcatus in the American Museum of Natural His-
tory and in the Museum of Comparative Zoology revealed a dif-
ference in the shape of the ear opening that initially was pre-
sumed to be a dimorphism : the ear opening was in one case
circular (Fig. 3), and in the other markedly elongate (Fig. 2).
Later, field work in Camaguey, Cuba, indicated that this was not
a phenomenon of polymorphism in a single species but instead
that two distinct species of the carolinensis group existed in
central Cuba.

Dr. Richard Etheridge independently discovered the two
conditions of the ear opening while studying the carolinensis
group at the University of Michigan Museum of Zoology. He

184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

called to our attention the apparent identity of the elongate-
eared Cuban form with Anolis allisoni Barbour (1928) from
Roatan Island off the coast of Honduras.

A direct comparison of the types of A. allisoni (M.C.Z. 26725,
26727-55) with specimens of both Cuban species in the Museum
of Comparative Zoology confirmed Dr. Etheridge's observations.
It was thus established that Anolis allisoni Barbour was an
available name for one of the two Cuban forms. It was neces-
sary, however, to determine the proper allocation of the much
older name Anolis porcatus Gray 1840. We therefore provided
Mr. J. C. Battersby of the British Museum (Natural History)
with material to compare with Gray's types. He very kindly
informed us that the name porcatus Gray applies to a round-
eared form.

A. porcatus has been customarily considered as merely one of
the subspecies of Caroline ns is (Barbour, 1937 ; Oliver, 19-18). For
the purposes of this paper and to facilitate discussion we refer
to the Cuban round-eared form as A. porcatus. The subspecies
of carolinensis have been described without any reference to the
pronounced geographic variation found within porcatus in Cuba.
Certainly the level of differentiation of some of the Bahaman
subspecies could be duplicated within the Cuban populations of
porcatus. However, subspeciation within porcatus and the rela-
tionship of the Cuban populations to the non-Cuban subspecies
of carolinensis is a taxonomic problem that need not concern us
here.

DEFINITION OF THE ANOLIS CAROLINENSIS GROUP

Long-snouted lizards having the nostril median to the canthal
ridge and separated from the rostral by three scales (Fig. 1) ; a
rostral that is bordered on the posterior dorsal margin by five
scales ; loreal rows usually 3-4 ; the anterior sublabials are wider
than long, supradigital scales multicarinate ; ventral scales at
midbody in transverse and diagonal rows; ventrals and dorsals
slightly keeled ; dorsals and laterals subequal in size, smaller

RUIBAL AND WILLIAMS: ANOLIS ALLISONI

185

than or equal to the ventrals; tail round in cross section. The
body color is variable and changeable ; it may be yellow, green,
grey, blue, brown or variegated. There is pronounced sexual
dimorphism. The males are larger, and are characterized by
well-marked frontal and/or canthal ridges. The head scales are
rugose; there is a reddish or mauve dewlap and enlarged post-
cloacal scales. In contrast, the females are smaller and lack the
enlarged postcloacal scales; they also lack frontal ridges or
elevated canthal ridges, the head scales are multicarinate, the
dewlap is small.

Figure 1. Dorsal view of the tip of the snout of a specimen of A. por-
catus from Oriente. The earolinensis group characters of the rostral are
shown: five scales bordering the rostral posteriorly and three scales be-
tween the rostral and nostril.

186

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

CANTHAL RIDGE

Figure 2. Head of a male A. allisoni from Camaguey. The elongate
ear opening, the high canthal ridges, and the small postorbital scales are
shown. The canthal ridges are so high that the frontal ridges are not
visible in a lateral view.

FRONTAL RIDGE

CANTHAL RIDGE

Figure 3. Head of a male A. porcatus from Camaguey. The circular
ear opening, the high frontal ridges, and the large postorbital scales arc
shown.

Figure 4. Head of a male A. allisoni from Roatan Island. The charac-
teristic overlapping rostral scale of this population is shown.

RUIBAL AND WILLIAMS: ANOLIS ALLISONI

187

COMPARISON OF ALLISONI AND PORCATUS

The following characters serve
porcatus :

A. allisoni (Fig. 2)

1. Ear opening is elongate, the pos- 1.
terior margin forming a long
longitudinal depression.

2. Temporal or postocular scales 2.
smaller.

3. Males with canthal ridges higher 3.
than the frontal ridges.

to distinguish allisoni from

A. porcatus (Fig. 3)
Ear opening circular, or oval, the
posterior margin not forming an
elongate depression.
Temporal or postocular scales
larger.

Males with the frontal ridges
higher than the canthal ridges.

The elongate ear opening is a distinctive character of allisoni
and is very different from the round or vertically oval ear open-
ing of the other species of Anolis. In allisoni the tympanum
resides in a depression having a sharply demarcated anterior
margin formed by the temporal scales. Posterior to the tym-
panum is a triangular, elongate, scaleless groove that tapers
posteriorly. Dorsally, the tympanum and groove are bounded
by a loose fold of skin (Fig. 5).

In the entire genus Anolis the only examples that we have seen
of an ear opening approximating this condition occur in some
specimens of porcatus from Pinar del Rio, Cuba. In these speci-
mens there is a short V-shaped posterior margin to the ear
(Fig. 6). This condition in porcatus is not identical to the elon-
gate depression in allisoni, but nevertheless does represent an
approximation. About half of the 65 specimens examined of
Pinar del Rio porcatus show some evidence of such a V-shaped
posterior margin.

3C«SS&

Figure 5. Ear opening of A. allisoni from Camaguey.

Figure 6. Ear opening of A. porcatus from Pinar del Eio. Some speci-
mens of the allopatric populations of porcatus from extreme western Cuba
show evidence of convergence toward allisoni, in having the posterior mar-
gin of the ear V-shaped and resembling the elongated posterior depression
of the allisoni ear.

188 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

J n life, the males of the two species are readily distinguishable
since allisoni males have a brilliant blue head and thorax which
is never seen in porcatus (Fig. 10). Gundlach (1880) in his
discussion of porcatus mentions the blue form as a variant of
porcatus. Similarly, Barbour and Ramsden in the "Herpetology
of Cuba" (1919) confused the two species and provided a color
description of porcatus that is a mingling of the characters of
both species. They noted that the species (actually allisoni) was
very common in the vicinity of Camaguey, and admit to having
examined specimens from all six provinces.

The blue and green color of the allisoni males can change to
an overall brown color. There is another transient color phase
in which the green part of the body and limbs assumes a yellow
color while the blue portion becomes a light violet color. The
throat of the males usually shows some blue pigment, particu-
larly in the area of the folded dewlap. A dark elongate spot
is usually evident above the forelimb. The males also have a
well-defined stripe from below the eye to the ear. The females
of Cuban allisoni show no blue color and are solid green, and like
the males can change to an overall brown. Living specimens of
allisoni from the Cuban provinces of Las Villas, Oriente, and
Camaguey do not differ in color or pattern. However, after
preservation with formalin most of the Las Villas specimens
show dark reticular markings over the nape and dorsum. These
black reticular markings are present in life only during the
brown color phase in the Camaguey specimens and are custom-
arily not seen in the preserved animals.1

The blue color so distinctive of allisoni males is never present
in porcatus.

VARIATION IN PORCATUS

A. porcatus shows pronounced geographic variation in color
pattern (see Table 1). In the western portion of Cuba (Pinar
del Rio and Habana provinces), porcatus can change in color
from green to brown and shows a pattern of dark (black or dark
green) reticulations over at least part of the body. There is also
an elongate dark patch above the forelimb and single scattered
white scales on the nape and parts of the body. Figure 7 shows

i We have seen two male allisoni which lacked the blue on the heart and trunk.
These animals were observed alive in the held (at the same locality) in Cam-
aguey for a number of days. One of the specimens was collected and preserved
(M.C.Z. 60928).

RUIBAL AND WILLIAMS: ANOLIS ALLISON1

189

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this type of pattern clearly. In the specimens from Oriente, at
the extreme eastern end of the island, the color change ranges
from green and grey-green to brown. There is a distinctive mid-
dorsal light stripe bordered by darker pigment. The chin and
throat area is well marked with longitudinal dark stripes. There
is an ocellus (a spherical or elongate dark spot surrounded by
white) above the forelimb. There is some evidence of reticular
body markings on some specimens and often scattered white
scales. Figure 8 is an illustration of an Oriente male of porcatus.

Figure 7. Male A. porcatus from Pinar del Rio. An example of a
' ' western ' ' porcatus.

In contrast to the eastern and western patterns the Camaguey
populations of porcatus in the center of the island have an over-
all color of grey or light green or brown. The larger males are
usually grey and some demonstrate no green color; however,
females and some males do demonstrate a light green phase.
Superimposed upon the grey color is a complicated pattern
(Fig. 9) of dark and light reticulations. The body as well as
the nape is covered with numerous white scales. The Camaguey

RUIBAL AND WILLIAMS: ANOLIS ALLISON]

191

porcatus are thus sharply distinguishable from the sympatric
and more abundant blue allisoni. The Camaguey specimens of
porcatus are further distinguished by the absence of a white
stripe below the eye which is characteristic and prominent in
allisoni. However, it is interesting that Pinar del Rio porcatus

Figure 8. Dorsolateral view of a male A. porcatus from Guantanamo,
Oriente. An example of an "eastern" porcatus.

show convergence to allisoni in having a white stripe below the
eye similar to that of allisoni. Some of the other characters that
readily distinguish the sympatric Camaguey porcatus from alli-
soni are not found outside the range of allisoni. The large flat
temporals of Camaguey porcatus, which differ markedly from
the small temporals of allisoni, are not so characteristic of the

192

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOOV

Oriente porcatus where only the postoculars appear to be en-
larged. Similarly some of the large males of porcatus from
Pinar del Rio have eanthal crests that are higher than the
frontal crests and thus further resemble allisoni in this charac-
ter. The specimens of porcatus from Las Villas (where allisoni
is also found) seem to resemble the Pinar del Rio porcatus in the
pattern of reticulations and in not being grey in color as is the
Camaguey porcatus. However, we have only seen preserved
Las Villas porcatus and therefore, do not know definitely whether
the specimens from there ever assume a grey color. We have
seen no live specimens from Matanzas province. The specimens
of porcatus from Habana Province resemble the Pinar del Rio
specimens.

The female specimens of porcatus do not demonstrate as
pronounced a geographic variation as the males. The female

Figure 9. Male A. porcatus from Camaguey. An example of a "cesntral"
porcatus.

Camaguey porcatus usually show well-marked white scales on
the lateral surfaces of the body and are thus readily dis-
tinguished from the eastern and western forms. The eastern
females, like the males, show a middorsal light stripe bordered
by a darker pigment. The females from western Cuba often
show a brownish middorsal stripe similar, but not identical, to
that of the eastern form. (Actually the middorsal stripe is a
common characteristic of the female of many species of Anolis.)
Even some female specimens of allisoni will at times have a
narrow middorsal brown stripe that contrasts with the overall

green.

RUIBAL AND WILLIAMS: ANOLIS ALLISONI

19:5

BLUE

GREEN

BLUE

GREEN

Figure 10. Male A. allisoni from Camaguey. The more heavily stippled
areas on the head and thoracic region are blue while the rest of
the body is green.

VARIATION IN ALLISONI

Anolis allisoni was originally described from Roatan Island by
Barbour (1928) who recognized that it was closely related to
porcatus and cited three scale characters by which it was sup-
posed to differ from that species . Not one of these characters
appears to be valid. His description of the male color pattern
agrees with that given above for the Cuban allisoni. However,
Barbour did not notice the peculiarity of the ear opening nor
did he recognize that the Roatan allisoni was the same species
that he had undoubtedly observed many times in Las Villas (at
Soledad) and Camaguey. We have examined the types and
paratypes in the Museum of Comparative Zoology, and a large
series of specimens of allisoni from the Chicago Natural History
Museum, and we can find onlv two characters that distinguish

104 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

the Islas de la Bahia specimens (from the Islands of Bonaea and
Koatan) from the Cuban specimens of allisoni. In the Islas de la
Bahia populations most of the males have an enlarged overlap-
ping rostral (see Fig. 4). These specimens, when viewed ven-
trally, show a markedly projecting rostral that overhangs the
tip of the lower jaw. Of a total of 34 male specimens with un-
damaged snouts, 24 (or 70%) had an overhanging rostral. In
contrast, the rostral in the Cuban specimens is usually only
slightly projecting. All of the Bonaea and Roatan specimens
show dark reticular markings over the dorsum and nape, and
also show white scales on the nape. This pattern resembles
that seen in the brown phase of allisoni and also is much like the
reticular pattern present on porcatus — from Pinar del Rio. In
all other characters studied, the Cuban and Islas de la Bahia
populations appear to be identical.

The distribution of allisoni is not limited to the Islas de la
Bahia and Cuba. Specimens of allisoni have also been collected
in Half Moon Cay (Schmidt, 1941), about 100 miles to the
northwest of Roatan, near Turneffe Island (C.N.H.M. 30541 and
34628-9 and M.C.Z.' 60983 — a total of 12 specimens). These
specimens lack the overhanging rostral of the Islas de la Bahia
forms. The color, in life, of the Half Moon Cay specimens is
not known. As preserved, the specimens have irregular blotches
of blue and purple and would thus appear to possess the blue
pigment characteristic of the Cuban and Islas de la Bahia popu-
lations. The absence or presence of blue after preservation is,
of course, not a definite indication of the true color in life.
Unfortunately, the color of anoles is subject to unpredictable
variation following death and preservation. Thus, the C.N.H.M.
specimens from Bonaea and Roatan are all brown with dark
reticulations and a single specimen from Half Moon Cay has the
same pattern. However, Barbour's description of the Roatan
populations does not mention such a pattern except as appearing
after preservation. It is therefore possible that the reticulated
pattern is made evident after preservation, and in life is only
evident when the animal is in the brown color phase. The Half
Moon Cay specimens, besides lacking an enlarged rostral, are
also distinctive in that no dark shoulder patch is evident on any
of the twelve specimens, that blue pigment is present on the
ventral surface of the body, and that the white head stripe while
evident below the orbit fades in the temporal area and is not
visible posterior to the ear opening.

RUIBAL AND WILLIAMS : ANOLIS ALLISON I

I!).",

Morphologically, allisoni is thus divisible into three groups

Cuba, Islas de la Bahia, and Half Moon Cay

CUBA

1. Rostral slightly
overlapping

2. Dark patch above
forelimb

3. White stripe from
ventral border of
orbit to ear and some
evidence of stripe
posterior to the ear

ISLAS DE LA BAHIA

1 . Rostral strongly
overlapping

2. Same as Cuban
population

3. Same as Cuban
population

HALF MOON CAY

1 . Rostral not
overlapping

2. No dark patch above
forelimb

3. White stripe on
ventral border of
orbit, fading in
temporal area, and
not evident posterior
to the ear

4. Belly color whitish

4. Same as Cuban
population

Belly color possibly
blue or green. Blue
in preserved speci-
mens, but color in
life unknown

No white scales on
nape

White scales on nape

No white scales on
nape

It is immediately evident from the above that the Cuban and
lslas de la Bahia populations are very similar while the Half
Moon Cay population stands out as distinct from the other two.
We believe that the morphological differentiation that has oc-
curred in the Half Moon Cay and Islas de Bahia populations is
evidence for assuming that these Central American populations
of allisoni are old in these islands and not recent introductions
by man. These three populations should probably be given sub-
specific designation to emphasize this fact. However, we have
refrained from this step in the hope that we may be able to
observe and compare living specimens of all three populations
and thus confirm the color differences.

The differentiation of the Half Moon Cay and Islas de la
Bahia populations is also evidence for assuming that the two
populations represent separate invasions from Cuba. Thus, the
Half Moon Cay population being the most differentiated can be
considered to be the oldest or first invaders, while the Islas de la
Bahia forms represent a more recent dispersal to the Central

19G BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

American shore and thus more resemble the Cuban population.
Certainly the differentiation of the two offshore populations
precludes our considering them as derived one from the other.

DISTRIBUTION

The round-eared form (or forms) which we are calling A.
porcatus is found throughout the island of Cuba and on Isla
de Pinos. At the western end of the island (Pinar del Rio and
Habana) and at the eastern end (Oriente) it is the very common
arboreal lizard found in well lighted areas around houses, in
gardens, in pastures, in the edges of the forests, and probably
in the forests in the higher sunnier portions of the trees. Next
to A. sagrei it is the most common lizard on the island. How-
ever, in the provinces of Camaguey and Las Villas, porcatus is
a rarer lizard. In this area allisoni is the common species, found
in the identical habitats that porcatus occupies in the eastern
and western portions of the island.

The abundance of allisoni in central Cuba, of porcatus in the
eastern and western portions of the island, and of sagrei through-
out the island is probably a recent phenomenon associated with
the destruction of the forest and the introduction of agriculture.
In a recent article (Ruibal, 1961) it was shown that allisoni
and sagrei are ecologically restricted to open areas of high
insolation. Both these species (and presumably porcatus as well)
demonstrate a high temperature preference and bask in open
exposed habitats. Previous to the destruction of the forests
these species were probably restricted to the savannas, the open
coastal vegetation (thickets), and similar sparsely covered areas.
Using Waibel's (1943) estimates about the former plant life of
Cuba, it appears reasonable to assume that probably 80 per cent
of pre-Columbian Cuba was covered with broadleaf or hardwood
forest. Currently only 11 per cent of the surface area of Cuba
remains covered with broadleaf forests (Smith, 1954). Allisoni,
porcatus, and sagrei thus represent species that originally were
restricted in their distribution in Cuba, but with the destruction
of the natural vegetation have spread and become the predom-
inant anoles of the island.

Figure 11 shows that the range of allisoni is completely in-
cluded within the range of porcatus. In Camaguey and Las

RIIUAL AND WILLIAMS: ANOLTS ALLISONI

197

o
&

I!'"1 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Villas the two species have not been collected together at all
localities; however, wherever intensive collecting of allisoni has
been done (Soledad, Trinidad, 9 km. west of Camaguey, and 15
km. southwest of Camaguey) specimens of porcatus have also
been collected. Mr. Kevin W. Marx of the University of Min-
nesota observed these two species in the vicinity of Soledad,
Las Villas, and was also impressed by the fact that the forms
are sympatric and that porcatus was the rarer species (personal
communication). Therefore, we believe that porcatus may be
found throughout the range of allisoni but as a less common
species. During field work in Camaguey in the summer of 1957
and 1959, only 12 specimens of porcatus were seen and collected
while the number of allisoni seen was certainly in the hundreds
— this, in spite of the fact that a purposeful search was made
for porcatus. The males of the Camaguey porcatus are, of
course, immediately recognized in the field due to their varie-
gated pattern and white scales. The females are also recognizable
but are not as brilliantly marked. Allisoni and porcatus in cen-
tral Cuba are sympatric species. The exact ecological interrela-
tionship of These two species is not known; nevertheless, the
available evidence indicates that their respective ecological niches
overlap. Both species have been collected in the same localities
and in the same situations : on fence posts, on buildings, corrals,
in banana groves, on palms (Coccothrinax) in coastal thickets,
and along the edges of forests. In contrast allisoni has often
been observed on the royal palm (Roystonea) and the coconut
palm while we have never observed Camaguey porcatus on these
palms. Similarly we have twice collected female porcatus inside
the forest, and have never observed allisoni in such a habitat.
In April of 1960 the senior author spent ten days in Camaguey
and observed numerous specimens of porcatus on the trunks
and branches of the "algarrobas" (Samanea saman) in a pasture
near Camaguey. A diligent search was made for allisoni in this
portion of the pasture and none was found. During these ten
days in April more specimens of porcatus were observed than
in the previous two summers of field work. It is thus possible
that allisoni and porcatus may demonstrate some form of eco-
logical temporal or seasonal replacement such as Neill and Allen
(1959) have claimed for some lizards from British Honduras.
However, the ecological data that we have obtained for porcatus
in Camaguey has been of a fortuitous nature and thus prevents
us from reaching any definite conclusions at this time.

RUIBAL AND WILLIAMS: ANOLIS ALLISONI 19!)

The distribution of the two species has been plotted on the
map in Figure 11 and the localities are listed at the end of the
text. These localities are those represented by specimens ex-
amined from the collections of the Museum of Comparative
Zoology, American Museum of Natural History, University of
Michigan Museum of Zoology, United States National Museum,
and the Chicago Natural History Museum. A. allisoni has so
far been recorded from Las Villas, Camaguey, and the lowland
western portion of Oriente. In contrast, porcatus is recorded
from all of the six provinces and Isla de Pinos. In Oriente the
distribution of porcatus is limited to the more mountainous
areas and there is actually a distributional gap between the
Oriente porcatus and the porcatus from Camaguey. This is a
gap of about 150 kilometers between the easternmost Camaguey
localities and the Oriente sites for porcatus. It is impossible at
this time to determine whether this is an actual break in the
range of porcatus or whether it merely reflects the lack of
collecting in lowland western Oriente. Similarly only three
specimens of porcatus all from one locality have been seen by
us from Matanzas province. This is clearly the consequence of
Matanzas being an area that has been singularly ignored by
herpetological collectors. Future collecting in Matanzas should
show porcatus to be as abundant as it is in Habana province.

Outside of Cuba, allisoni is found on Bonaca and Roatan in
the Islas de la Bahia and at Half Moon Cay near Turneffe
Island. It would appear probable that it is distributed through-
out the other small islands in the Gulf of Honduras and it may
possibly also be found in the neighboring mainland. A. allisoni
thus has a disjunct distribution. The Little Caymans located
part way between Cuba and the coast of Honduras are populated
by A. maynardi, a form round-eared like porcatus and not close
to allisoni.

The distribution of allisoni from central Cuba to the offshore
islands of Honduras shows the most distant dispersal of any
of the carolinensis group. It immediately raises the question
of the evolutionary origin of allisoni. Any attempt to explain
the zoogeography of allisoni and porcatus requires an analysis
of the evolution and zoogeography of the entire carolinensis
group. It would appear best to leave this aspect of the discussion
to a later paper.

2(J() BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

DISCUSSION

In all of the preceding discussion, the assumption has been
that we are dealing with two species, one of which (allisoni)
has a central distribution in Cuba while the second (porcatus)
is represented by three geographic races spread along the entire
length of the island. We made this assumption to facilitate the
presentation of the data, and it is now necessary to critically
examine the distribution and morphology of the Cuban forms
and determine what other interpretations may be made of the
data.

The variations described above for allisoni and porcatus per-
mit us to distinguish, in Cuba, four morphological forms of the
carolinensis group: allisoni, "western" porcatus, "central" por-
catus, and "eastern" porcatus. Using these names now as simple
labels without any taxonomic implications will allow us to
analyze the distributional patterns. From the field observations
in Camaguey and the morphology of allisoni and "central"
porcatus it is obvious that these two sympatric forms are sep-
arate species. No morphological intermediates between allisoni
and porcatus are known from Camaguey and Las Villas prov-
inces. However, the relationship of these two central species
to the eastern and western forms is not immediately apparent.
At least the following four alternative interpretations (see Table
2) are possible:

Hypothesis 1. The "western" porcatus and allisoni are geo-
graphic races of one species, and the "central" and "eastern"
porcatus are geographic races of another species. The main
evidence in support of this hypothesis is that only in the "west-
ern" porcatus do we find an elongate ear opening approximat-
ing the external ear opening of allisoni. Furthermore, some of
the large "western" porcatus males resemble allisoni in having
a large canthal ridge. However, this hypothesis must assume
a zone of morphological intergradation between the Pinar del
Rio "western" porcatus and the allisoni from Las Villas and
Camaguey. This hypothesis is untenable since the specimens
from Habana and the few from Matanzas show no evidence of
being morphologically intermediate between allisoni and por-
catus. In addition, the "central" porcatus show a color pattern
that closely resembles the variegated pattern of "western" por-
catus. The hypothesis further assumes that the "central" and

eastern" porcatus are geographic races and should therefore

i i

BUIBAL AND WILLIAMS: AXOLIS ALLISONI 201

demonstrate morphological intermediates between the two races
in Oriente. As mentioned previously, there is a distributional
gap between the "central" par cat us in Camaguey and the "east-
ern" porcatus in Oriente (see Fig. 11). This gap may be more
apparent than real ; nevertheless, on present evidence it is a
distributional hiatus.

Hypothesis 2. This is the reverse of the preceding hypothesis.
In this instance, allisoni is considered to be a geographic race
of "eastern" porcatus while the "central" and "western"
porcatus would be geographic races of a second species. The
similarity in ear and canthal ridges between allisoni and "west-
ern" porcatus, previously mentioned, would be interpreted as
the result of convergence. Thus in central Cuba where the
two species overlap they demonstrate pronounced divergence
(allisoni as contrasted to "central" porcatus) while in the zones
where a single species is to be found there appears to be mor-
phological convergence ("eastern" and "western" porcatus).
This interpretation would consider the carolincnsis group in
Cuba as an example of "character displacement" similar to
previously cited cases of this type (Brown and Wilson, 1956).
In support of this hypothesis it can be said that the patterns of
the "central" and "western" porcatus are similar and can be
easily imagined to be derived one from the other. The specimens
of porcatus from Ilabana and Matanzas are variegated and do
not present an obstacle to this interpretation as they do to the
first hypothesis.

A further corollary of this hypothesis is that intermediate
populations between allisoni and "eastern" porcatus should be
found in Oriente. There is support for this from four male
specimens (U.S.N.M. 138117, 138126-28) collected by Mr. Jerry
D. Hard}', Jr., about 10 miles north of Calio Cruz and at nearby
Jucural. All specimens resemble allisoni but have the following
porcatus-like characters :

1. A round ear opening in one specimen and the others with
only a shallow groove posterior to the ear opening.

2. Two of the specimens show white spots on the nuchal area.

3. Three of the specimens demonstrate postoculars and tem-
porals that appear to be intermediate in size between allisoni
and porcatus.

However, a single female from the same area (U.S.N.M. 138125
from Jucural) shows no evidence of porcatus characters.

202 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

A series of 18 specimens from Cabo Cruz collected by Albert
Schwartz also show evidence of hybridization between allisoni
and "eastern" porcatus. Of the 11 males in the series only one
shows a well developed ear depression while the others have
poorly developed shallow areas posterior to the ear opening.
These ears are morphologically very similar to some of the Pinar
del Rio porcatus (Fig. 6). One male also has white spots on the
nape. Four of the females have well developed allisoni-\ike ear
openings. A single male (with an allisoni ear) shows a light
middorsal stripe bordered by darker pigment like that of "east-
ern" porcatus.

The area of the hybrids (Fig. 11) lies where one would assume
a zone of contact between allisoni and porcatus; allisoni is in
the lowland western portions of Oriente while porcatus is in the
more mountainous southern and eastern portions. Porcatus is
found throughout the Sierra Maestra and would be expected to
contact allisoni on the northern slopes and along the coastal
area near Cabo Cruz. Collecting in this area indicates that the
intermediate zone between allisoni and "eastern" porcatus must
in any case be narrow — at San Ramon, 20 ml. northeast of this
area "typical" allisoni have been collected.

Hypothesis 3. Allisoni is a species limited to the central por-
tion of the island while porcatus is islandwide in its distribution.
(The "eastern," "central" and "western" porcatus represent
geographic races of the same species.) This, of course, is the
way in which the data were presented preceding this section.
We may now consider the evidence for and against this theory :

a) "Central" porcatus and allisoni are sympatric and do
not interbreed. This fact at least establishes the distinctness of
the two forms in Las Villas and Camaguey.

b) "Central" porcatus, though different from "eastern" and
"western" porcatus in color pattern, shows many similarities
to these two forms (structure of the ear opening, temporals,
reticular markings, white scales, and low canthal ridges).

c) "Eastern" and "western" porcatus morphologically re-
semble each other more than either of them does allisoni.

Therefore, on morphological grounds alone it would be pos-
sible to consider "central" porcatus conspecific with the "east-
ern" and "western" forms. The "central" porcatus would
still represent an example of character displacement where the

RUIBAL AND WILLIAMS: ANOLIS ALLISONI 203

species is sympatric with allisoni. The "eastern" and "west-
ern" populations would in this case be considered to be con-
nected across the length of Cuba by the rarer "central" porcatus.

This interpretation is weakened by the apparent distribution
gap mentioned previously between "central" and "eastern"
porcatus. The gap may or may not be real and all that we can
do is hope that intensive collecting can be done in this area in
the near future. It should be pointed out that a similar distribu-
tional gap exists in this area for Anolis allogus and A. homo-
lechis (Ruibal and Williams, 1961). Furthermore, a comparable
situation exists at the western end of the distribution of allisoni,
in the province of Matanzas where very few records are avail-
able for any of the Cuban species of Anolis. Both the eastern
and western distributional gaps may merely reflect the lack of
herpetological collecting that has been done in these areas.

Also weakening this interpretation is the presence of hybrids
in the area of Cabo Cruz, Oriente, between "eastern" porcatus
and allisoni. However, the hybrids do not necessarily commit
us to discarding this hypothesis. Sufficient cases are known in
various animal species (Blair, 1941; Volpe, 1959; Gilliard, 1959)
where hybrids occur when the ecological isolation between
species has been disturbed. The evidence for intermediates be-
tween allisoni and "eastern" porcatus indicates that the zone of
contact between the two morphological types must be narrow
and that it corresponds to the border between the lowland
agricultural areas and the less modified mountains. The Cabo
Cruz area may represent a situation where the isolating mech-
anisms between the two species have broken down in an eco-
logically disturbed zone (i.e., where agriculture is actively en-
croaching into a forested area). It is of course also possible
that a narrow zone of hybridization between allisoni and por-
catus may exist throughout the periphery of the distribution of
allisoni. In central Cuba the two species are presumed to have
evolved mechanisms to reduce interspecific competition. The
existence of character displacement and the fact that in the area
of sympatry porcatus is less common and occupies a more re-
stricted habitat than where it is allopatric to allisoni suggest
that isolating mechanisms (behavioral, ecological, etc.) have
evolved. However, at the periphery of the distribution of
allisoni this species would be adjacent to populations of por-
catus that have not been subjected to competition with allisoni.
These populations of porcatus would be occupying the habitats

204 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

in which allisoni has proved superior (such as the arboreal
habitat, around houses and gardens). If these populations are
brought into contact, by let us say a human disturbance of the
habitat or by an expansion of one of the populations, an initial
hybridization between the species is not unreasonable. With
time, selection would operate so that each species would be
segregated in the ecological niche to which it was best adapted
and the hybrids would be eliminated. This of course presup-
poses that the hybrids are not as successful or well adapted as the
parental species. Unfortunately, we have no data concerning the
viability of the hybrids from Cabo Cruz or the precise ecology
of this area.

Hypothesis 4. The final hypothesis to be considered is a three-
speeies one (see Table 2). In this case allisoni and the "eastern"
porcatus would be two separate species while the "western" and
•"central" porcatus would constitute a third species. This
assumes that the distributional gap in Oriente between "cen-
tral" porcatus and "eastern" porcatus is a real one or at least
not occupied by populations morphologically intermediate be-
tween "central" and "eastern" porcatus. It would also assume
that the hybrids between allisoni and "eastern" porcatus are
explainable in the same manner as for the previous hypothesis.

Three of these hypotheses can be discarded if collecting in the
distributional gap between "central" and "eastern" porcatus
yields specimens that are morphological intermediates between
these two forms. The only hypothesis that would be retained
would then be the third case presented — allisoni as a centrally
distributed species with porcatus represented by three geographi-
cal races, "eastern," "central" and "western." If no inter-
mediates are found in this zone then a more detailed analysis
of the zone of contact between allisoni and porcatus in Oriente
will be mandatory.

RUIBAL AND WILLIAMS: ANOLIS ALLISONI

205

TABLE 2. Diagram of the four hypotheses that are
considered applicable to the data presented on A. allisoni and A. porcatus .

WESTERN CUBA

CENTRAL CUBA

EASTERN CUBA

[western porcatus

- subspecies-

HYPOTHESIS I
(2 species)

allisoni]

(central porcatus

- subsneci?3 — — > eastern pifcatusl

HYPOTHESIS 2
(2 species)

[western porcatus <—

[allisoni <-—

-subspecies > central porcatus]

- subspecies — -^ eastern porcatus!

HYPOTHESIS 3
(2 species)

[western porcatus

[allisoni]

-subspecies ^central porcatus <-

■ subspecies :^> eastern porcatus!

HYPOTHESIS 4
(3 species)

[western porcatus <z-

(allisoni!

-subspecies 5. central porcatus!

[eastern porcatus]

SUMMARY

1. The carolinensis group of AnoUs is defined and two closely
related Cuban representatives of the group, porcatus and allisoni
are described.

2. A. allisoni is found in central Cuba (Las Villas, Cama-
guey, and western Oriente) and is sympatric with porcatus in
this area. A. porcatus is apparently island-wide in its distribu-
tion and is a common species in eastern and westernmost Cuba,
but it is rare in central Cuba.

3. A. allisoni is also found on a number of islands off the
Central American mainland. Evidence is presented that indi-
cates that allisoni is not a recent introduction by man into these
offshore islands.

4. On Cuba, allisoni and porcatus show evidence of character
displacement and convergence.

5. Both species on Cuba are considered examples of species
that were preadapted to occupy the areas cleared of forest and
modified by man's activities.

6. Three morphological forms of porcatus are described in
Cuba, ''western" porcatus, a "central" porcatus, and an "east-
ern" porcatus.

7. Evidence of hybridization between allisoni and "eastern"
porcatus is presented.

8. Four hypotheses arc presented to explain the distribution
and biology of the three forms of porcatus and of allisoni.

206 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ACKNOWLEDGMENTS

We are indebted to Sr. Ramon Molina of Caniaguey for first
showing us the variegated A. porcatus of central Cuba. Dr.
Albert Schwartz of Albright College generously provided us
with data and specimens which permitted us to better evaluate
these species. We wish to thank Dr. Richard Etheridge of the
University of Michigan and Mr. Stanley Rand of Harvard Uni-
versity for their advice and interest in the problem. We are
particularly indebted to Mrs. Theresa McKnight and Mr. John
Healy of the staff of the American Museum of Natural History
for their enthusiastic and interminable assistance in regard to
the loan of specimens. To Charles Bogert of the American
Museum of Natural History, Robert Inger of the Chicago Natu-
ral History Museum, and Doris Cochran of the United States
National Museum, we are grateful for the loan of specimens.
Miss Dereth Bogert made the excellent drawings that accom-
pany the text.

This investigation is part of a study of Cuban anoles financed
by National Science Foundation Grant No. G-5634.

LOCALITY LIST
Hybrids

Cuba

Orient e: Cabo Cruz; 10 ml. N. of Cabo Cruz; Jucural (be-
tween Cabo Cruz and Niquero).

A. porcatus

Cuba

Pinar del Rio: San Diego de los Bahos; Vihales; Guane;
Pinar del Rio; San Vicente; just N. of San Vicente; near Isabel
Rubio; Cayo la Reina near Puerto Esperanza; nr. Herradura;
Guanajay; Dimas.

Habana: Habana; Regla; San Antonio de los Bahos ; San
Jose de las Lajas; Isla de Pinos.

Matanzas: Alacranes (Alfonso XII).

Las Villas: Rodas; Bahos de Ciego Montero ; Soledad; Central
Pnrio; Caibarien; Sierra de Trinidad; Trinidad; Cienfuegos.

Camaguey: Marti; 9 km. W. of Camaguey; 15 km. S.W. of
Camaguey; Playa Sta. Lucia (E. of the Bahia de Nuevitas) ;
Sierra de Cubitas ; Sierra de Najasa; Loma de Cunagua.

RUIBAL AND WILLIAMS: AXOLIS ALLISONI 207

Orient e : Santiago; Jiguani; Upper Rio Ovando ; Coast S. of
Pico Turquino ; Pico Turquino : Imias ; Banes ; Baracoa ; Guan-
tanamo : Buey Arriba ; Sagua de Tanamo ; Moa ; Miranda ; 27
km. S. of Yara; mouth of Rio Yumuri; Cananova; Calabazar;
Bayate.

A. allisoni

Cuba

Las Villas: San Isabel de la Lajas; Banos de Ciego Montero ;
Trinidad; Soledad; Caibarien; Central Covadonga; Rodas; San
Jose del Lago ; Central Caracas.

Camaguey: Camaguey; Marti; Cascorro; 15 km. S.W. of
Camaguey; Playa Sta. Lucia (B. of the Bahia de Nuevitas) ; 13-
20 km. S. of Playa Sta, Lucia; 7 km. N.E. of Sta. Cruz del Sur;
Sta. Cruz del Sur; 9 km. W. of Camaguey; Bahia de Nuevitas;
Moron; Banao ; Tana.

Oriente: Birama; Omaja; San Ramon; 5 ml. S.W. of Man-
zanillo.

Honduras

Islas de la Bahia: Roatan; Bonaca.

British Honduras
Half Moon Cay

LITERATURE CITED

Barbour, T.

1928. Reptiles from the Bay Islands. Proe. New Eng. Zool. Club 10:
55-61.

1937. Third list of Antillean reptiles and amphibians. Bull. Mus.
Comp. Zool. 82: 17-166.
Barbour, T. and C. T. Ramsden

1919. The herpetology of Cuba. Mem. Mus. Comp. Zool. 47: 71-213.
Blair, A. P.

1941. Variation, isolation mechanisms, and hybridization in certain
toads. Genetics 26: 398-417.
Brown, W. L. and E. 0. Wilson

1956. Character displacement. Syst. Zool. 5: 49-64.
Gilliard, E. T.

1959. The ecology of hybridization in New Guinea honey eaters (Aves).
Amer. Mus. Nov. no. 1937: 1-26.

208 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Gray, J. E.

1840. Catalogue of the species of reptiles collected in Cuba by W. S.
MacLeay, Esq., Ann. Mag. Nat. Hist., 5: 108-115.

GUNDLACH, J.

1880. Contribucion a la erpetologia cubana. G. Montiel, Habana 1-98.
Xeill, W. T. and R. Allen

1959. Studies on the amphibians and reptiles of British Honduras.
Publ. Res. Div. Ross Allen Rept. Inst., 2, no. 1 : 1-76.
Oliver, J.

1948. The anoline lizards of Bimini, Bahamas. Amer. Mus. Nov.
no. 1383: 1-36.
Ruibai,, R.

1961. Thermal relations of five species of tropical lizards. Evolution,
15: 98-111.
Ruiral, R. and E. E. Williams

1961. The taxonomy of the Anolis homolechis complex of Cuba. Bull.
Mus. Comp. Zool. 125: 209-246.
Schmidt, K. P.

1941. The amphibians and reptiles of British Honduras. Zool. Ser.
Field Mus. 22: 475-510.
Smith, E. E.

1954. The forests of Cuba. Maria Moors Cabot Found., no. 2: 1-98.
Volpe, E. P.

1959. Experimental and natural hybridization between Bufo terrestris
and Bufo foivleri. Amer. Midland Nat. 61 : 395-412.
Waibel, L.

1943. Place names as an aid in the reconstruction of the original vege-
tation of Cuba. Geogr. Rev. 33: 376-396.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 125, No. 8

THE TAXONOMY OF THE ANOLIS HOMOLECHIS
COMPLEX OP CUBA

By
rodolfo ruibal

Division of Life Sciences
University of California, Riverside

and

Ernest E. Williams

Museum of Comparative Zoology

Harvard University

CAMBRIDGE, MASS., U.S.A.

printed for the museum
August, 1961

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Bulletin of the Museum of Comparative Zoology

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Vol. 125, No. S

THE TAXONOMY OF THE ANOLIS HOMOLECHIS
COMPLEX OP CUBA

By
rodolfo ruibal

Division of Life Sciences
University of California, Biverside

and

Ernest E. Williams

Museum of Comparative Zoology

Harvard University

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

August, 1961

No. 8 — The Taxonomy of the Anolis homolechis Complex of

Cuba,

By Rodolfo Ruibal and Ernest E. Williams

INTRODUCTION

In Barbour and Ramsden's (1919) "Herpetology of Cuba"
nineteen species of the genus Anolis were recognized. Of these,
five were grouped together as related, in both the key to the spe-
cies and in the text. These species are: A. homolechis (Cope)
1864, A. mestrei Barbour and Ranisden 1916, A. rubribarbus Bar-
bour and Ramsden 1919, A. quadriocellifer Barbour and Rams-
den 1919, and A. allogus Barbour and Ramsden 1919. Later, two
more species were described that were related to this group :
A. ahli Barbour 1925, and A. patricius Barbour 1929. In 1937
Barbour published the "Third list of Anglican reptiles and
amphibians. ' ' At this time he further emphasized the interrela-
tion of the various species in the group by making trinomials of
all names, as follows :

A. allogus allogus

A. allogus mestrei

A. allogus ahli

A. homolechis homolechis

A. homolechis rubribarbus

A. homolechis quadriocellifer

A. homolechis patricius
It must be pointed out that A. mestrei was described in 1916,
three years before A. allogus. Consequently, the trinomials in-
volving these two names are incorrect as a matter of nomencla-
ture. Furthermore, a careful analysis of the morphology of these
six described forms indicates that Barbour was also wrong in
regard to the species and subspecies groupings of the various
forms.

In the sections that follow an attempt is made to redescribe
the forms on a sound morphological basis and to determine the
interrelationship of the various species. The data for this work
have been obtained by a study of the types of five of the six de-
scribed forms (only the type of homolechis was not seen), and
the specimens of these species in the Museum of Comparative

212 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Zoology and the American Museum of Natural History, as well
as the paratypes of A. mestrei from the United States National
Museum in Washington. Further information was obtained
during six weeks of field work in 1957 in the Province of
Camaguey, Cuba, as well as ten weeks of field work in 1959 in
Camaguey, Oriente, Las Villas, and Pinar del Rio.

THE HOMOLECHIS GROUP

All of the named forms here discussed can be distinguished
from the other Cuban species of A?wlis by the following com-
plex of characters: Head scales usually keeled, with a single
median keel or with two or more keels; ventral scales smooth,
with a convex posterior border, and in diagonal and/or longi-
tudinal rows; dorsals small, non-imbricate, smooth or keeled,
the middorsal scales slightly larger but grading in size imper-
ceptibly to the smaller laterals ; tail laterally compressed. Frontal
ridges and a sharp canthus rostralis. Marked sexual dimor-
phism. The females are smaller, have multikeeled head scales,
the dewlap is absent or very small, and they have a different color
pattern. The males are larger, have usually a single keel to the
head scales, have a large dewlap, and often have a well-developed
fin to the tail.

The species of this group are all forest-dwelling forms. They
are customarily to be found in shaded areas but some occur in
the sunnier margins of the forest.

6 '

CHARACTERS

Preserved specimens of the various species that compose the
liomolcchis complex are not readily distinguishable. Scale
characters can be used to separate most of the species; however,
all of the scale characters are subject to considerable variation.
In the field, color and pattern differences allow immediate and
absolute identification of the species but these are usually de-
stroyed or obliterated during the fixation and preservation of
the animals. In the field also, the species distinctions are made
more evident by the ecological separation observable between
the various forms. The unravelling of the various described
species in this complex has been possible only because the field
work in Cuba allowed us to observe the lizards while alive.

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 213

The most useful diagnostic characters are listed and defined
below:1

1. The number of scales that separate the supraorbital semi-
circles. There may be from 0-3 scales between the semicircles.

2. The type of suture between the mentals and the small post-
liiental scales. This may be a transverse suture (Figure 6b) or
the small postmentals may separate the posterior medial margins
of the mentals (Figure 6a).

3. The structure of the supracarpal and supradigital scales.
These scales may be smooth, have a single keel, or be multi-
carinate and mucronate (Figure 7).

4. The number of scales between the first canthals. This may
vary from 4-12 scales. The first canthal is defined as the an-
teriormost enlarged and elongate canthal scale.

5. The structure of the scale anterior to the external naris.
This scale may be single or divided by a median horizontal
suture (Figure 8).

6. The size of the scales along the posterior border of the
interparietal. These scales may be large and sharply distinguish-
able from the dorsals (Figure 3) or they may be small and
grade into the dorsals (Figure 4).

7. The head/ear ratio as an indication of the size of the ear.
The head length is measured with Vernier calipers and is the
distance from the tip of the snout to the anterior border of the ear

EAR HEIGHT

HEAD LENGTH

Figure 1. The head length and ear height measurements. These dis-
tances are best measured with the use of calipers.

i The scale terminology used is that defined by Smith (1946).

214

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

opening (Figure 1). The ear height is measured with the cal-
ipers and is the maximum distance from the ventral to the
dorsal edge of the ear opening (Figure 1).

8. The body/femoral ratio as an indication of the length of the
hind limbs. The body length is the snout-vent length measured
with a ruler and is the distance from the tip of the snout to the
vent. The femoral length is taken with a ruler and with the
femoral region perpendicular to the body, and is the distance
from the midventral line (pubic symphysis) to the knee (Fig-
ure 2).

FEMORAL LENGTH

Figure 2. The femoral length measurement,
reference is the pubic symphysis.

The midventral point of

9. The color and pattern of the animal. The dewlap color and
pattern easily distinguish the species in the field but these
striking differences usually disappear or are little evident in
preserved specimens. The body color and pattern are also very
useful in the field ; these are sometimes still evident in preserved
specimens.

RUIBAL AND WILLIAMS: ANOLIS HOMOLECHIS COMPLEX 215

THE SPECIES

A definition of each of the species is provided below as well
as general comments on the taxonomy, distribution, and ecology
of the forms. Unless there is a statement to the contrary, all
descriptions and comments are based exclusively on male speci-
mens. In general the females of the various species can be
distinguished by the same scale characters that distinguish the
males, and differ also in the color pattern of the body. Except
in one case all color descriptions in this paper are based on live
specimens.

With the exception of ahli, all the species of the homolechis
group may have high caudal fins in the males. This, however,
is a variable character and often, within a single population,
some males are devoid of a caudal crest while others have the
crest well developed. The few (12) males of ahli that we have
seen lacked a caudal fin.

The list of the localities for the species is given at the end
of the text. The data were obtained from the specimens in the
American Museum of Natural History, the Museum of Compara-
tive Zoology, and the United States National Museum.

Anolis allogts Barbour and Ramsden

A. allogus Barbour and Bamsden, 1919.
A. allogus allogus, Barbour, 1937.
A. abatus Ahl, 1924.

Type locality. Bueycito, S. of Bayamo, Oriente, Cuba.

Definition. Supraorbital semicircles separated by two scales
(Figures 4 and 9); a transverse suture between the mental and
postmentals (Figure 6b) ; supracarpal and supradigital scales
multicarinate and mucronate (Figure 7b) ; usually with 7-!)
scales between the first canthals (Figure 10) ; scale anterior to
the naris divided by a horizontal suture (Figure 8) ; scales
around the posterior margin of the interparietal small and
grading into dorsals (Figure 4). The body/femoral ratio aver-
aging 3.3 (Table 1) ; the head/ear ratio averaging 6.2 (Table 2).
Maximum snout-to-vent length : S , 58 mm., 9 , 41 mm.

Body color of $ I in life with reddish and yellow reticula-
tions. Color may change from a pale pink or tan to a very
dark red-brown. Yellow reticulations usually prominent when
in dark color phase. Tail is usually redder than the body.
Iris metallic blue or grey. Usually some evidence of four dark

216

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

chevrons on dorsum. Dewlap ground color ranging from light
yellow to apricot with 3-4 reddish stripes and a white margin
(Figure 11).

The females have a wide middorsal light stripe (tan or yellow
in color) while the dorso-lateral areas are a darker brown color.
The middorsal stripe may have a scalloped margin and in a few
specimens evidence of diamond-like markings in the light stripe
may be seen.

Figure 3. Dorsal view of the head of Anolis homolechis.
Figure 4. Dorsal view of the head of Anolis allogus.

Taxonomy. We have examined the type specimen of Anolis
abatus Ahl and regard it as synonymous with allogus.

Our description of allogus differs from the color description
provided by Barbour and Ramsden. They describe the body
color as brown to black and the dewlap as yellow with a carmine
spot. To resolve this discrepancy a collecting trip was under-
taken to the type locality in the northern foothills of the Sierra
Maestra. There is no longer any forest in the vicinity of Buey-
cito and it was necessary to travel farther into the foothills to
Buey Arriba (8 km. south of Bueycito) to find forest and collect
the species. The specimens from Buey Arriba have a yellow -tan

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 217

dewlap with three bright red stripes, and the body color is
reddish as described above for the species. We feel that the
color description of Barbour and Ramsden is probably errone-
ous. The red stripes in allogus are not sharply demarcated and

Figure 5. Dorsal view of the head of Anolis rnestrei.

Figure 6. Ventral view of the chin showing the diagnostic postmental
scales, (a) A. Iiomolechis ; (b) A. allogus.

218 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

often are very near to each other. In alcoholic specimens the
three or four separate stripes often appear confluent. In all
other respects the type specimen and the other specimens that
Barbour ascribed to allogus are in agreement with our descrip-
tion.

Distribution. This species is recorded from all of the provinces
except Matanzas and Las Villas ; it is also absent from the Isla
de Pinos (see Figure 13). The lack of records from Matanzas
and from the portion of Las Villas outside the Sierra de Trini-
dad may be the consequence of insufficient collecting in these
areas, or agriculture may have destro}^ed the natural habitat of
the species.

Variation. There is no evidence of any significant geographic
variation. The color and pattern of the dewlap is variable (see
Figure 11) but we do not consider the variants well enough
defined to merit taxonomic recognition. At Buey Arriba in
Oriente the two or three red stripes are bright red, while in
many of the Camaguej- specimens the stripes are often very
faint and of an orange color. Specimens may show from two to
four stripes. In Pinar del Rio the background color is apricot
while in most of the other localities it is best described as
yellowish or tan.

Some of the specimens from Oriente Province (Baracoa, coast
south of Pico Turquino, mountains north of Imias) though not
distinguishable from allogus by any scale characters, show a
general pigmentation pattern that differs from that of other
Oriente allogus. As preserved, these specimens are devoid of a
body pattern and show a pale bod}7 coloration (tan) and a dark
dewlap. It is, of course, not improbable that the mountainous
areas of Oriente will prove to have forms related to allogus that
are distinguishable only by the dewlap or body color (as are ahli
and rubribarbus). Isolation of populations in the various valleys
and ranges of Oriente would permit the evolution of distinct
forms. However, the detection and recognition of such forms
requires detailed data from the living animals.

Ecology. The ecology of this species is discussed in some
detail in Buibal (1061). It is a forest-dwelling species restricted
to the cooler, deeply-shaded portions of the broadleaf forests,
perching on small tree trunks a few feet from the ground.
Characteristically not a shy species, it will usually allow the col-
lector to approach within a foot before escaping by running
down to the base of the trunk or to the ground.

KIJIBAL AND WILLIAMS: ANOLIS IIOMOLECHIS COMPLEX 219

220 bulletin : museum of comparative zoology

Table 1. The Body/Femoral Ratio of the Six Species

The values for homolechis and allogus include samples from Pinar
del Rio, Camaguey and Oriente. The range for each species
is in parentheses below the mean.

Species X Mean a aM

homolechis 169 3.6 .16 .01

allogus 96 3.3 .21 .02

ahli 11 3.3 .16 .05

Mean

L69

3.6

(3.1-4.0)

3.3

(3.0-3.8)

3.3

(3.0-3.6)

3.3

(2.7-3.6)

3.2

(2.9-3.5)

3.1

rubribarbm 17 3.3 .15 .04

mestrei 24 3.2 .15 .03

nn i as

Table 2. The Head/Ear Ratio of the Slx Species

The values for homolechis and allogus include samples from

Pinar del Rio, Camaguey and Oriente. The range for each

species is in parentheses below the mean.

Species

Mean

homolechis

162

7.7
(6.0-9.7)

.45

.04

allogus

100

6.2

(4.9-8.2)

.70

.10

ahli

6.3

(5.7-6.8)

.38

.11

rubribarbus

7.6
(6.3-8.6)

.75

.18

mestrei

8.5
(6.2-9.7)

.87

.20

imias 1 6.0

RUIBAL AND WILLIAMS: ANOLIS HOMOLECHIS COMPLEX 221

The females appear to be more terrestrial than the males, and
are often found in the leaf litter on the floor of the forest and
at the base of the small trees.

This species is absent from the coastal forests, savannas, pine
forests, and from agricultural areas. We have never observed it
near human habitations or in gardens.

Previous to the destruction of the Cuban forest, allogus was
probably a widespread and common species. With the destruc-
tion of its natural habitat this species is now restricted to the
small patches of dense forest that remain on the island. It has
been collected at many localities in the Sierra Maestra and as
high as 4000-6000 feet on Loma Cordero near Pico Turquino.

The apparent absence of allogus from the Isla de Pinos (where
homolechis is found) may be the result of the specialized ecologi-
cal requirements of this species for deeply-shaded, cool forests.
The forests of Isla de Pinos may not be dense enough to provide
this type of habitat. Furthermore, over-water invasion of the
Isla de Pinos is hampered by the low thermal tolerance of
allogus.

Anolis ahli Barbour

A. ahli Barbour, 1925.

A. allogus ahli, Barbour, 1937.

Definition. The scalation of this species cannot be dis-
tinguished from that of A. allogus. The body/femoral ratio and
head/ear ratios are also identical to allogus (see Tables 1 and
2). The maximum snout-to-vent length: $, 58 mm. and 9,
43 mm.

The body color of this species usually shows a greenish cast
and the general color may shift from tan to dark brown. A
common body pattern is a "salt and pepper" speckling. Usually
four dark saddle markings are visible on the dorsum. The iris
is blue. The dewlap has a large red spot that is surrounded by
a yellow-white area (Figure 11).

Taxonomy. In the 1937 checklist, Barbour placed this form
under allogus as a subspecies. The two forms are allopatric;
the nearest recorded locality for allogus is at the Loma de
Cunagua in western Camaguey. We have been conservative in
utilizing the trinomial and prefer to view ahli as a full species.
The lack of scale character differences between ahli and allogus
does not prevent their being considered full species: the body

222 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

color and dewlap color pattern sharply distinguish the two
forms. Color patterns are certainly as "important" as scalation
in distinguishing species.

Distribution. This species is known only from the Sierra de
Trinidad in Las Villas.

Ecology. This species has been collected in the darkest, shaded
parts of the forest in the Sierra de Trinidad. It perches a few
feet from the ground on tree trunks. It is a shy species and
usualty will run to the base of the trunk or to the ground on
the approach of a collector. It is often difficult to distinguish
the lizards from the background in the dim light of the forest.
It appears to be the ecological equivalent of allogus in the Sierra
de Trinidad.

Hardy (1958) has described the use of the tail by this species
as a prehensile organ. "We have observed that all of the species
of the homolcchis complex can curl the tail laterally into a tight
whorl. We have, however, failed to observe any actual pre-
hensile use of the tail. As mentioned previously, this species
is the only one of the homolechis group in which no males have
been observed with well developed caudal crests.

Anolis rubribarbus Barbour and Ramsden

A. rubribarbus Barbour and Eamsden, 1919.
A. homoledhis rubribarbus, Barbour, 1937.

Type locality. Puerto de Cananova ; near Sagua de Tanamo,
Oriente, Cuba.

Definition. The scalation of this species is the same as that of
allogus except for the structure of the scale anterior to the naris.
In most specimens of rubribarbus this is a single scale that
extends from the margin of the nares to the rostral (Figure 8).

ROSTRAL

HOMOLECHIS ALLOGUS

Figure 8. The structure of the scale anterior to the external naris in
A. homolechis and A. allogus. In A. rubribarbus the scale is usually single
as in A. homolechis.

RUIBAL AND WILLIAMS: ANOLIS HOMOLECHIS COMPLEX 223

In allogus and ahli this scale is divided by a horizontal suture.
The body /femoral ratio averages 3.3 and is identical to that of
allogus (Table 1). The ear is smaller than that of allogus or
ahli and the head/ear ratio average is 7.6, very close to that of
homolcchis (Table 2). The maximum snout-to-vent length: $,
62 mm., 2 , 42 mm.

The general body color of this species is usually grey. It
ranges from a pale grey to an almost black ground color. Yellow
spots and reticulations may be present on the sides. The body
may show no pattern with only a pale grey color or a "salt and
pepper" marking. However, the most characteristic color phase
is a pattern of dark (blackish) vertical bands separated by
lighter (yellowish or grey) bands (see Figure 14). In this
phase there are about six vertical bands on the body and more
on the tail. The limbs are also banded. The dark body bands
are usually narrower in the middorsal area and widen laterally,
while the lighter bands show the reverse. The iris is blue-grey.
This species has the most brilliant and distinctive dewlap pat-
tern of any of the Cuban anoles (Figure 11)- — 4-5 thin red
stripes on a deep yellow ground. The scales along the edge of
the dewlap are large and white and provide a white margin to
the dewlap. The scales on the inner portions of the dewlap are
smaller and black. The chin has well-marked dark reticular
markings.

We have no color data on the female specimens of rubribarbus.

Taxonomy. This species was erroneously considered a sub-
species of homolechis by Barbour. Rubribarbus is actually closely
related to allogus, and museum specimens are difficult to dis-
tinguish from allogus-. the undivided character of the scale
anterior to the nares serves to separate most specimens of rubri-
barbus when color is absent. This species is further distinguish-
able from allogus by the smaller ear opening. However, there is
considerable overlap in the latter character and it is therefore
not very useful in practice. Some color characters, however,
assist in identifying preserved specimens, e.g., the presence of
darkly pigmented scales at the base of the dewlap of rubribar-
bus. This usually appears as a dark area in the throat of
specimens. In allogus usually no dark pigment is visible in the
throat. In those specimens of allogus that demonstrate a dark
throat the pigment is between the scales rather than in the scales
as in rubribarbus. Some specimens of rubribarbus also show

224

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

homolechis

87*

78%-

allogus

0 12 3 4

Figure 9. Histograms of the number of scales between the supra-
orbital semicircles of allogus and homolechis. The great majority of the
specimens of these two species are distinguishable by this character.

50-

Ld
O

i 25-

-z.

o
cr

homolechis

allogus

4 5 6 7 8 9 10 II 12
NUMBER OF SCALES BETWEEN CANTHALS

Figure 10. Histograms of the number of scales between the canthals
in homolechis and allogus. There is considerable overlap between the
specimens of these two species.

RUIBAL AND WILLIAMS : ANOLIS IIOMOLECHIS COMPLEX 225

REDDISH

allogus

YELLOW
WHITE

allogus

ahli

RED

YELLOW

mestrei

YELLOW-ORANGE

RED

WHITE

rubribarbus

Figure 11. Diagrams of the dewlap pattern and color of A. allogus,
ahli, rubribarbus, and mestrei.

226

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

indications of the dark vertical body stripes which are not
present in allogus. Also of value in distinguishing rvbribarbus
are the well-marked reticulations on the chin.

In their description (Barbour and Ramsden, 1919), the
authors listed M.C.Z. 11941 as the number of the type specimen
and gave M.C.Z. 11868 as the number of the paratype (and of
the specimen figured in plate 9). There is an evident lapsus
since M.C.Z. 11868 is the actual type specimen, while the number
11941 belongs to a specimen of Platysaurns capensis from Rho-
desia.

YELLOW

WHITE

homolechis

WHITE

YELLOW

homolechis

WHITE

homolechis

WHIT

REY

homolechis

''WJS&j^f

h. quadriocellifer

Figure 12. Diagrams of the dewlap pattern and color of A. homolechis.

RUIBAL AND WILLIAMS: ANOLIS HOMOLECHIS COMPLEX 227

J-

• r-l

-<J

-Q

•rH

f-i

-M

•r-H

«w

cc1

•c*

CO
iH

S-.

.f-t

t-a

228

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Distribution. This species is allopatric to allogus. Bubribar-
bus is so far known only from the northeast coast of Oriente,
from Cananova to Punta Gorda to the east of Moa (Figure 13).

Ecology. We have collected this species to the east of Moa in
the broad leaf gallery forests that extend along the streams and

Figure 14. Diagram of the distinctive dark banding of A. rubribarbus.

rivers that descend through the coastal pine forests. We have
also collected it in the cooler broad-leaf forests southeast of
Moa at an elevation of about 1000 feet. It is a forest-dwelling
species but does not appear to be restricted to the deep shaded
portions of the forest as are allogus and ahli. It is a shy species
and difficult to observe against the greyish bark of some of the
trees. It perches head down a few feet from the ground in the
same fashion as the other species.

It is very probable that this species may no longer exist in
the vicinity of Cananova, the type locality. We visited this
locality briefly in 1959 and were unable to find forests to collect
in.

Anolis iiomolechis (Cope)

Xiphosurus homolechis Cope, 1864.

Anolis homolechis, Boulenger, 188,"); Barbour, 1914; Barbour and Ramsden,

1919.
A. oalliurus Aid, 1924.
A. muelleri Ahl, 1924.
A. cubanus Ahl, 1925.
A. patrioius Barbour, 1929.
A. homolechis homolechis Barbour, 1937.
A. homolechis patricius Barbour, 1937.
A. quadriocellifer Barbour and Ramsden, 1919.

Type locality: "West Indies.'* It would be reasonable to re-
strict the type locality to Habana, Habana Province, Cuba. The

RUIBAL AND WILLIAMS : ANOLIS HOMOLECIIIS COMPLEX 229

populations in the vicinity of the city of Habana have a pure
white dewlap.

Definition. Supraorbital semicircles separated by a single scale
(Figures 3 and 9) ; posterior medial margins of the mentals
separated by small postmentals (Figure 6a) ; supracarpal and
supradigital scales usually smooth or with a single keel (Figure
7a) ; usually 5-7 scales between the first canthals (Figures 3 and
10) ; a single undivided scale anterior to the nares and in contact
with the rostral (Figure 8) ; scales along the posterior margin
of the interparietal large and sharply demarcated from the

Figure 15. Dorsal pattern of the female specimens of A. homolechis. In
life the pattern is composed of black and various shades of brown.

dorsals (Figure 3). The body /femoral ratio averages 3.6; the
head/ear ratio averages 7.7 (Tables 1 and 2). The maximum
snout-to-vent length: $ , 56 mm., 9 , 43 mm.

230

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

The general body color ranges from a very light tan, through
brown, to black. Usually there is some evidence of horizontal
stripes on the lateral surface of the body, and of four dark
chevrons on the dorsum. Yellow markings may be present
laterally. The iris is gold or metallic brown in color. The dew-
lap color of this species is very variable (Figure 12) and is
discussed below in detail.

The female color pattern usually consists of a series of light
colored diamonds on the dorsum (Figure 15). The general color
may change from tan to black. The ventral surface is sometimes
yellow in females.

Taxonomy. We have examined the types of the three species
described by Ahl {calliurus, muelleri, and cubanus) and con-
sider them synonymous with homolecliis. We have also examined
the type of A. patricius Barbour from Mina Piloto in Oriente
and find no character to distinguish this form from homolecliis.
The type locality of patricius is in the municipality of Sagua
de Tanamo but we have been unable to locate Mina Piloto pre-
cisely. No information about the dewlap color was provided in
the description of the type. The homolecliis from the town of
Sagua de Tanamo and nearby Cananova have a white dewlap.

In the 1937 checklist, Barbour included A. quadriocellifer
from the Ensenada de Cajon, Cabo San Antonio, at the extreme
western end of Cuba, as a subspecies of homolecliis. We have
examined the types as well as additional specimens from the
type locality collected by Albert Schwartz. This form is readily
distinguishable from the other populations of homolecliis by the
light -margined, dark ocellus above the foreleg (Figure 16), and

Figure 16. The lateral pattern of a male specimen of A. homolecliis
quadriocellifer (M.C.Z. 11907) from Ensenada de Cajon, Pinar del Eio.

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 231

the yellow dewlap with three reddish stripes (Figure 12). The
juvenile as well as female specimens of quadriocellifcr have a
well-marked lateral ocellus. We agree with Barbour in consider-
ing this form a subspecies of homolechis. We have reached this
decision from a consideration of the morphology of the popula-
tions near the Ensenada de Corrientes as represented by speci-
mens collected by Albert Schwartz and his staff. (The localities
are shown on map of Figure 17.)

The Ensenada de Corrientes is midway between Cabo San
Antonio, the type area for quadriocellifer, and the towns of
Cayuco and Isabel Rubio (formerly Mendoza) in the vicinity
of which typical white dewlap homolechis has been collected.

Specimens from Ensenada de Corrientes have a yellow dewlap
and have white spots on the sides of the body that resemble the
light-colored margins of the quadriocellifcr ocelli. Specimens of
quadriocellijer have most of the supracarpal scales with two or
three keels. Specimens from the Ensenada de Corrientes have
most of these scales with only one or two keels, and two of the
specimens (of a total of 23 males examined) have all the supra-
carpal scales smooth. Specimens from the vicinity of Cayuco
and Isabel Rubio have the supracarpal scales with only a
single keel or smooth.

On the basis of these characters we infer that typical quadrio-
cellifcr from Cabo San Antonio is connected to "typical" homo-
lechis from southern Pinar del Rio by an intermediate popula-
tion (only adult males used in the comparison) as shown in the
following table:

quadriocellifer

(8 specimens)

Populations from

Ensenada de Corriente:

(23 specimens)

homolechis

(14 specimens from

SW of Cayuco)

a) Yellow dewlap with a) Yellow dewlap,
red stripes.

b) White-margined
lateral ocellus.

a) White dewlap,
b) Lateral white spots. b) No white spots.

c) Supracarpals usually c) Supraearpals usually c) Supracarpals usually
with 2-3 keels. with 1-2 keels. with 0-1 keel.

Four of the specimens (representing the three localities listed
in the above chart) were unique for homolechis in having the
ventrals with slight keels.

232 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Variation. No geographical variation was found in the scala-
tion of this species. However, there is a pronounced variation
in the color and pattern of the dewlap. The majority of the
populations of homolechis throughout the island and the Isla de
Pinos have a white dewlap. The dewlap may be pure white
showing only a faint indication of black pigment or there may
be two or three grey stripes on a white background (Figure
12). Both these types show no systematic geographical distribu-
tion and may actually be found in the same population. The
populations showing these two white patterns may best be
referred to as the "white dewlap" form. Another group of
variations may be called the "yellow dewlap" form. We have
personally seen this form from the Sierra de Cubitas, the shores
of the Bahia de Nuevitas, and along the north coast at the Playa
Santa Lucia (east of the Bahia de Nuevitas) all in the Province
of Camaguey. The Sierra de Cubitas population has a yellow
dewlap with a broad white margin (Figure 12). The popula-
tions from the vicinity of the Bahia de Nuevitas and Playa
Santa Lucia have a deeper yellow or orange color, a narrow
white margin, and one or two stripes of white or light yellow
(Figure 12). The two yellow dewlap populations are therefore
distinguishable. We do not know if these two populations are
isolated from each other or not. However, it appears probable
that they are separated by the savanna that extends north to the
coast near the Rio Maximo.

P. J. Darlington has also recorded yellow (and/or orange)
dewlap homolechis from the south coast of Oriente near Pico
Turquino, Cabo Maisi, and the lower Rio Ovando. Albert
Schwartz has also collected the yellow dewlap form along the
south coast of Oriente from just north of Cabo Cruz to Playa
-Juragua, east of Siboney. Yellow dewlap forms have also been
recorded at Banes on the north shore of Oriente, and of course
the previously mentioned population from the Ensenada de
Corrientes in Pinar del Rio has a yellow dewlap. We cannot
compare the color patterns of these populations with those of
the Camaguey yellow dewlap populations because we have not
seen the former in life, and beyond the fact that they are yellow
we do not know the details of the pattern.

The map (Figure 17) indicates the known distribution of the
white and yellow dewlapped forms. There is an apparent gap
in the distribution of the coastal yellow dewlap in southern
Oriente between Cabo Maisi and Playa Juragua. In this zone

RUIBAL AND WILLIAMS: ANOLIS HOMOLECHIS COMPLEX 233

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234 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

white dewlapped forms have been collected. Similarly, there is
an apparent gap on the north coast of Oriente. As mentioned
above there is also a gap between the yellow forms of the Sierra
de Cubitas and the yellow forms of Playa Santa Lucia in
Camaguey. For the moment we believe it prudent to refrain
from designating these populations as subspecies or species. With
the exception of the Sierra de Cubitas population all of the
other yellow forms are coastal. The coastal forests are usually
drier and warmer than the more inland forest and the yellow
forms may represent a homolechis ecotype adapted to the more
stringent conditions of the coastal areas. Collecting in the
coastal forest in other parts of the island may prove that the
yellow form is more widespread than the present data indicate.
We have studied the contact of the yellow and white forms
near the Playa Santa Lucia, Camaguey. The senior author
undertook field work in this area in 1957 (Paiibal, 1958) and
both of us visited the area in 1959. A road runs inland from
the beach at Sta. Lucia in a southwesterly direction. Collec-
tions were made at various stations along this road from the
beach to 21 kilometers inland. At the shore the vegetation is a
coastal thicket predominantly made up of seagrape (Coccoloba)
and a small palm (Coccothrinax sp.). A few kilometers inland,
broadleaf forest is found on limestone. Some of the forest is in
relatively good condition, having only been ' ' highgraded ; ' ' other
parts have been severely cut for charcoal. Some tongues of man-
grove extend into the forest. At about 20 kilometers inland there
is only a sparse open forest with very few large trees. Cattle are
grazed in the area and most of the vegetation here is "mije"
(Eugenia), an arborescent cactus (Dcndrocerens) , and numer-
ous species of palms. A total of four visits were made to Santa
Lucia to sample the populations. From the thicket on the shore
to 12 km. inland only yellow homolechis were collected. From
13 km. to 17 km. inland, about 25 specimens of the white dewlap
form have been collected, yet within this same area (at the 15
km. and 17 km. stations) two specimens of the yellow form have
also been found. From the 18 km. to 21 km. stations only white
forms were found. This transition of yellow forms on the coast
and white forms further inland is similar to the situation that
P. J. Darlington found along the south coast of Oriente where
he collected both forms of homolechis, the vellow near the coast

BUIBAL AND WILLIAMS: ANOLIS HOMOLECHIS COMPLEX 235

and the white inland. No "intermediate" specimens between
the two forms have been found in the Sta. Lucia area.2 The
amount of field work done at Sta. Lucia was limited and we did
not obtain any ecological or behavioral data on the two forms.
As mentioned above, the yellow and white forms of homolechis
are best left, for the time being, without any taxonomic designa-
tion. It is to be hoped that in the near future a more precise
study can be made of this interesting problem.

Ecology. Some aspects of the ecology of this species are dis-
cussed in Ruibal (1961). In the province of Camaguey this
species is restricted to the margins of the broadleaf forests. It
is customarily found in areas of filtered sunlight — along paths,
small clearings, and the edges of the forests. However, it is found
throughout the drier and sparser coastal broadleaf forest where
allogus is absent. It is also found in the palm-pine savannas
in northern Oriente. In Camaguey this species is never found
near human habitations or in agricultural areas. In contrast, at
Sagua de Tanamo in Oriente, homolechis is an abundant lizard
of the fence posts around houses and pastures. It occupies the
same fence posts with A. sagrei and A. porcatus. Similarly, near
Habana we have observed homolechis in gardens. In Las Villas
we have had little experience with the species but we always
found it in forests or in the vicinity of forests. It is interesting
that the restriction of homolechis to forest habitats in central
Cuba may be correlated with the presence of A. allisoni (Ruibal
and Williams, 1961) around human habitations in central Cuba.
It may be that in eastern and western Cuba homolechis can
occupy the area around human habitations because allisoni is
absent (in these areas porcatus replaces allisoni [see Ruibal and
Williams, op. cit.]).

Male specimens of homolechis are characteristically found
perched head down on small tree trunks a few feet off the
ground. The tail is often curled laterally. The females are more
terrestrial and are usually on the ground or on perches closer
to the ground than the males.

This species is found throughout the forests of the Sierra Maes-
tra and has been collected as high as 5900 feet at Palma Mocha,
near Pico Turquino.

2 One specimen collected at 12 km. from the beach in 1957 had a red ground
color to the dewlap and yellow stripes.

236 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Anolis mestrei Barbour and Ramsden

A. mestrei Barbour and Ramsden, 1916.
A. allogus mestrei, Barbour, 1937.

Type locality. Valle de Luis Lazo, Pinar del Rio, Cuba.

Definition. Except for one character, we have been unable to
successfully distinguish the scalation of this species from that
of homolechis : Specimens of mestrei have small granular pos-
terior supraciliaries while most homolechis have larger, elongate,
and keeled posterior supraciliaries. This character is variable,
however, often subjective, and difficult to use. Mestrei further
differs from homolechis in having longer hind legs, the body/
femoral ratio averaging 3.2 (Table 1) and a smaller ear opening,
the head/ear ratio averaging 8.5 (Table 2). The maximum snout-
to-vent length $ , 55 mm., $ , 44 mm.

The body color of this species varies from dark to light grey
with an overall greenish cast. Yellow or orange spots are pres-
ent over the body. The iris is yellowish. The dewlap has a dark
red basal spot with two yellow-orange stripes. The remaining
broad margin is white (Figure 11). The scales on the dewlap
are white.

The females show the same general body color but can also
shift to a light brown color. There are darker hour-glass shaped
markings on the dorsum. The females have a small apricot
colored dewlap.

Taxonomy. An examination of the type and paratypes has
revealed that two species were confused in the original descrip-
tion. The type (M.C.Z. 11285) and paratypes (M.C.Z. 11286,
U.S.N.M. 26731 and 26733) are mestrei while two other para-
types (U.S.N.M. 26732 and 26344) are actually specimens of
allogus.

Barbour was in error in making mestrei a subspecies of
allogus in the 1937 checklist. Mestrei is readily distinguishable
from allogus by many scale and color differences. In the lime-
stone hills of Pinar del Rio the two species are sympatric.

Preserved specimens of mestrei are very difficult to distinguish
from homolechis. Usually there is a dark basal portion to the
dewlap in preserved specimens of mestrei. The superciliaries,
the length of the hind limbs, and the smaller ear opening will also
assist in distinguishing specimens. The ear opening in mestrei
is not only smaller (in height) but is also differently shaped
than in homolechis. In mestrei the opening is circular while in
homolechis it is higher than wide. This is readily apparent in

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 237

the comparison of the ear height/ear width ratio of the two
species :

homolechis mestrei

(15 specimens) (15 specimens)

Mean 1.59 1.03

Range 1.2-2.1 0.9-1.4

Distribution. This species is restricted to the broadleaf forests
of the limestone mountains and hills of Pinar del Rio — the
Sierra de los Organos and the Sierra del Rosario.

Ecology. We have observed this species in a forest in a small
"mogote" near Sumidero. Here mestrei was found throughout
the forested portion of the mogote. The females and juveniles
were on the ground or on the boulders of limestone that covered
much of the forest floor. The adult males appear to be re-
stricted to the limestone or were found on fallen logs near
the limestone outcrops. In no instance did we find the mestrei
perched head down on vertical tree trunks in the manner of
allogus and homolechis. The lizards would escape by running
to the ground and hiding, or by entering crevasses in the lime-
stone. "We started collecting at this locality at 8:00 in the
morning and during the early part of the morning only juveniles
and females were seen, and it was not until near noon that we
observed the adult males. This species appears to be restricted
to the shaded portions of the forest and only in a few instances
was it found in areas of filtered sunlight.

Anolis imias, sp. nov.

Type: M.C.Z. 42556, adult male, collected east of Guan-
tanamo Bay at Imias, on the south coast of Oriente Province,
Cuba, in August 1936, by P. J. Darlington.

Paratype. M.C.Z. 42555, adult female having the same data
as the type.

Diagnosis. Similar to A. homolechis but differing from that
species in having smooth brachial scales, smooth supraoculars,
larger ear opening, longer hind limbs, the gulars bordering the
mental along a transverse suture, and a brown dewlap.

Description of type. Head. Most of the head scales smooth,
the anterior-most scales with blunt keels. Six scales across the
snout between the first canthals. A frontal depression and
Aveakly developed frontal ridges. Nostril separated from the

238 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

rostral by two scales ou the right side and by one scale on
the left side. Seven scales bordering the rostral posteriorly.
Supraorbital semicircles separated from each other by a single
row of small scales. Supraoculars irregular in shape, smooth,
and separated from the supraorbitals by a row of scales. Canthus
well-marked. Four rows of loreals below the second canthal.
Suboculars keeled and in contact with the supralabials. The
subocular ring is continuous with a group of slightly enlarged
postoculars. Eight supralabials. A large interparietal with no
evidence of a parietal eye. Parietal region in a depression de-
marcated posteriorly by the V-shaped ridge of the underlying
parietal bone. Postparietal scales large and sharply demar-
cated from dorsals. Temporals small, the upper temporals
forming a longitudinal zone of scales larger than the granular
lower temporals and larger than the scales between the upper
temporals and the parietals. Ear opening vertical^ elongate.

Mental longitudinally divided, bordered along a straight
transverse contact by four small gulars. The mental is also
bordered posteriorly by a pair of sublabials and infralabials.
Throat and chin scales smooth. Dewlap large and with smooth
scales.

Body. Middorsal scales keeled and larger than the lateral
granular scales, but not sharply demarcated. Ventrals smooth,
imbricate, with a convex posterior margin, and in longitudinal
and diagonal rows.

Limbs. Humeral scales with weak keels, but the larger
brachial scales are smooth. Hind limb scales smooth with the
exception of the small scales on the dorsal surface of the limbs.
Most scales of the pes and maims smooth ; if keeled only with
a single weak keel.

Tail. Laterally compressed, with a high crest, and with all
the scales keeled. Verticils not readily distinguishable.

Measurements. Snout-to-vent, 65 mm.; head, 18 mm.; femoral
length, 21 mm.; car height, 3 mm. Body /femoral ratio = 3.1,
and the head/ear ratio = 6.0.

Description of the paratype. The female paratype resembles
the type in all respects except the following: all the head scales
keeled ; both nostrils separated from the rostral by a single
scale ; supraorbital semicircles separated by a double row of
small scales; rostral bordered posteriorly by six scales; mental
bordered posteriorly by only two small gulars along a straight
margin; tail slightly compressed but without a crest.

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 239

Measurements. Snout-to-vent, 46 mm.; head, 13 mm.; femur.
14 mm. ; height of ear, 2 mm.

Color. According to the collector, P. J. Darlington, the type
had a brown dewlap in life. As preserved, the only distinctive
markings are 12 vertical dark bands on the tail that are sep-
arated from each other by narrower lighter bands. Each hind
limb has six transverse dark bands, and the forelimb shows
evidence of about four transverse dark bands. The dorsum
(from the nape to the base of the tail) shows five indistinct,
dark crossbands. The chin has reticular markings.

The female paratype has the chin covered with dark reticula-
tions that are continuous with the dark vertical marks on the
labials. The body shows no discernible pattern.

Remarks. Though morphologically similar to homolcchis the
new species is a very distinctive form. The smooth head scales
and brachials set it off from all the other members of the homo-
lechis group. It resembles allogus in the postmental-mental
suture, the long hind limbs, and large ear opening.

P. J. Darlington also collected homolechis at Imias, but did not
collect any allogus. It is therefore possible that imias may be
the ecological equivalent of allogus.

DISCUSSION

Ecology. We have observed homolechis and allogus in broad-
leaf forest localities in Pinar del Rio, Camaguey and Oriente.
At all of these localities both species proved to have identical
perching habits (head down, a few feet from the ground, on
tree trunks) but were ecologically separated, allogus being re-
stricted to the deeper shaded portions of the forest while homo-
lcchis was found in the small clearings and paths or in the
sparser portions of the forest. In the forest habitat homolcchis
dwells in the filtered sunlight areas rather than in the deep shade
(where allogus is found) or in the open full sun areas (where
sagrei is found). This distinction between the species is reflected
in the mean body temperature of the species — homolechis having
a mean body temperature of 31.8°C. in contrast to 29.2°C for
allogus (Ruibal, 1961).

Our experience with ahli in the Sierra de Trinidad leads us
to believe that ahli resembles allogus in its ecology. In the Sierra
de Trinidad we succeeded in finding ahli only in the deeply

240 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

shaded portions of the forest. Our limited experience with rubri-
barbus in the vicinity of Moa indicated that this species may not
be as restricted to shade as allogus. Most of the specimens of
rubribarbus were collected in a portion of the hardwood gallery
forest that had been partially burned. The specimens of rubri-
barbus were seen on the exposed tree trunks in the clearings.

A. mestrei is apparently a shade-dwelling form like allogus.
However, it is distinguished from the other species in apparently
being restricted to limestone substratum, rather than to tree
trunks.

A. homolechis is characterized by a wider tolerance of habitats
than the other species mentioned above. It is not restricted to
the forests, and in eastern and western Cuba is found associated
with sagrei in the vicinity of human dwellings.

Distribution. Both maps (Figures 13 and 17) demonstrate
distributional gaps of homolechis and allogus in the region of
Matanzas and most of Las Villas. A similar gap occurs in N.W.
Oriente where no records of either species are known. We believe
that these are apparent gaps and that they are the consequence
of two factors :

1. Matanzas and Las Villas are intensively cultivated and the
natural forest habitat of these species has been almost com-
pletely obliterated. Allogus is nowhere known to survive out-
side of its shade-forest habitat and homolechis is only sometimes
found outside of the forest. Before the advent of agriculture
most of the vegetation of these provinces was hardwood forest
and it can be assumed that these two species were then common
and widespread in these areas.

'2. These areas are poorly collected and if appropriate habi-
tats still exist they have not been visited by herpetological col-
lectors.

It is of interest that similar distributional gaps exist for A.
porcatus and A. allisoni (Ruibal and Williams, 1961).

One question that further collecting in Las Villas may answer
is the relation of ahli to allogus. Our data so far indicate that
ahli is restricted to the Sierra de Trinidad; however, we have
no information whatever about the portions of Las Villas outside
of these mountains.

We lack similar information in respect to the zone of contact
between allogus and rubribarbus. If intermediates between these
two forms are found along the northern coast of Oriente it
would be necessary to reduce rubribarbus and allogus to sub-
species.

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 241

The distribution of the various species of this group can be
summarized in the following manner :

1. Islandwide distribution. The species sympatric, but eco-
logically isolated from each other :

homolechis and allogus

2. Local species inhabiting restricted areas and occupying an
ecological niche comparable to that of allogus. These species are
all allopatric to allogus:

alili, rubribarbus (?), and imias (?)

3. Local species inhabiting restricted areas and occupying
an ecological niche comparable to that of allogus, differing,
however, in the substratum selected for perching. Sympatric
with allogus-.

mestrei

Relationships. The six species of the homolechis group are
closely related, and as has been mentioned previously some of
the forms cannot be adequately distinguished by scale charac-
ters. The two most distinctive forms are allogus and homolechis.
The six species can be grouped in the following manner :

allogus j

ahli Very similar, not readily distinguishable except by color.

rubribarbus

homolechis
mestrei

Very similar, not readily distinguishable except by color.

imias Appears to be closer to homolechis than to allogus.

Another species which is closely related to these species, and
may actually merit being included in the group is A. sagrei. So
far the only distinguishing character of squamation that we have
been able to find to separate sagrei from the homolechis group
(especially homolechis itself) is the keeled mucronate condition
of the ventral scales in sagrei, and this keeling may sometimes
be very weak and even apparently absent (e.g. in some speci-
mens from Trinidad, Las Villas). Sagrei does, of course, differ
from homolechis and all other members of the homelechis group
in dewlap color and in thermal requirements and ecology.

In Table 3 the various characters used in distinguishing
species of the homolechis group are tabulated to facilitate a com-
parison of the six forms.

242

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

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244 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ACKNOWLEDGMENTS

We are indebted to Mr. Rudesindo Cuevas, the grandfather
of the senior author, and to Sr. Ramon Mousset for the hospi-
tality and assistance provided at the Finca Santa Teresa in
Camaguey. Sr. Ramon Molina was indispensable in providing
field assistance and in collecting material. Dr. Albert Schwartz
of Albright College was most generous and helpful in providing
information as well as specimens of the anoles that he has col-
lected in Cuba. We are grateful to Charles Bogert of the Ameri-
can Museum of Natural History, and Doris Cochran of the
United States National Museum for the loan of specimens in
their collections, and to Dr. Heinz Wermuth of the Berlin
Museum for allowing us to borrow the types of Aid's species.
Most of the drawings are the work of Miss Dereth Bogert.

This investigation is part of a study of Cuban anoles financed
by National Science Foundation Grant No. G-5634.

LIST OF LOCALITIES

A. allogus

PINAR DEL RIO : Sumidero; near Cabezas; 13.5 km. S. of Las
Pozas; Rangel; San Vicente; San Diego de los Banos; 8 km.
E. of Matahambre; Pinar del Rio; Soroa.

HABANA : 6.8 m. W. of Jaruco.

CAMAGUEY : 15 km. S.AV. of Camaguey ; nr. Banao, Sierra de
Cubitas; S. of Jaronu; Sierra de Najasa; 7 km. S.E. of Sta.
Cruz del Sur; Loma de Cunagua.

ORIENTE : Mai Paso, nr. Guantanamo ; Monte Libano, nr
Guantanamo ; Bueycito ; Los Negros, nr. Jiguani ; Baracoa
coast S. of Pico Turquino; Cobre Range, Sierra Maestra
Palma Mocha Mts., Sierra Maestra ; Banes ; Pico Turquino ; nr
Buey Arriba; 16 m. E. of Mayari ; Jutinicu ; mts. N. of Imias

A. ahli

LAS VILLAS : Electric plant, Sierra de Trinidad ; nr. Camana-
yagua, Sierra de Trinidad; W. slope of Sierra de Trinidad;
S. of Topes de Collantes ; Habanilla Falls, Sierra de Trinidad ;
4 km. W., 12 km. N. of Trinidad.

A. rubribarbus

ORIENTE: nr. Moa; Cananova; Mina Piloto.

A. mestrei

RUIBAL AND WILLIAMS : ANOLIS HOMOLECHIS COMPLEX 245

PINAR DEL RIO : San Vicente ; 10 km. W. of Cabezas ; 10 km.
N. of Cabezas; 2.9 km. B. of Isabel Rubio; Soroa; 8 km. E.
of Matahambre ; Rangel ; San Diego de los Banos ; Luis Lazo ;
Sumidero.

A. imias
ORIENTS: Imias.

Anolis homolechis homolechis
(Localities preceded by an asterisk denote yellow dewlap

populations)

PINAR DEL RIO : Luis Lazo ; Guane ; Sumidero ; nr. Consola-
cion del Sur ; San Diego de los Banos ; San Vicente ; N. of
San Vicente; nr. Cabezas; Soroa; 7.6 ml. E. Isabel Rubio;
2.9 ml. E. Isabel Rubio; 7-10 km. S.W. of Cayuco; *N. shore
Ensenada de Corrientes ; *W. coast Cabo Corrientes ; 8.5 ml.
E. Cabanas; San Cristobal; 1 m. N. of La Coloma; nr. Vinales.

HABANA : 9 km. S.W. San Jose de las Lajas ; Playa de Guanabo,
E. of Habana; Jibacoa; Isla de Pinos (various localities);
Iiabana, nr. Rancho Boyero ; Habana ; San Antonio de los
Banos ; Madruga.

MATANZAS: Pan de Matanzas; 6 km. N.E. of Matanzas; 5 km.
N.E. of Canasi.

LAS VILLAS : Topes de Collantes, Sierra de Trinidad ; Central
Soledad; Sierra de Jatibonico.

CAMAGUEY: 15 km. S.W. of Camaguey; Sierra de Najasa; 27
km. W. of Ciego de Avila; about 15 km. S.W. of Vertientes;
7-8 km. N.E. of Santa Cruz del Sur; *Sierra de Cubitas; *nr.
Banao ; *Bahia de Nuevitas, San Jacinto ; *Bahia de Nuevi-
tas, Los Ballenatos; *Loma de Cunagua, 12 m. E. of Moron;
*between Esmeralda and Jaronu ; *S. of Jaronu ; 0.6 ml. N. of
Majagua; Marti: Cuatro Caminos; *Playa Sta. Lucia and a
number of localities S.W. of Sta. Lucia.

ORIENTE : Guantanamo ; Sagua de Tanamo ; Cananova ; nr.
Moa; 16 km. E. of Mayari ; nr. Buey Arriba; Birama; Pico
Turquino; *Coast S. of Pico Turquino; Mina Piloto ; Buenos
Aires; near Santiago; *Banes; *Cabo Maisi; Los Negros,
nr. Jiguani ; Baracoa ; *lower Rio Ovando ; Sierra del Cobre ;
*Cabo Cruz ; N. of Imias ; Imias ; *Playa Juragua, nr. Siboney ;
*between Belie and Cabo Cruz.

A. homolechis quadriocellifer

PINAR DEL RIO: Ensenada de Cajon; Cabo San Antonio.

246 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

LITERATURE CITED

Ahl, E.

1924. Neue Iguaniden aus dem Zoologisehen Museum Berlin. Zool.
Anz. 62: 85-88.

1925. Neue Reptilien und Batrachier aus dem Zoologisehen Museum
Berlin. Archiv. f. Naturgesch. 90: 246-254.

Barbour, T.

1914. A contribution to the zoogeography of the West Indies, with
special reference to amphibians and reptiles. Mem. Mus. Comp.
Zool. 44: 209-346.

1925. A new Cuban Anolis. Occ. Paps. Boston Soe. Nat. Hist. 5:
167-168.

1929 Another new Cuban Anolis. Proc. New England Zool. Club 11:
37-38.

1937. Third list of Antillean reptiles and amphibians. Bull. Mus.
Comp. Zool. 82: 77-166.
Barbour, T. and C. T. Ramsden

1916. A new Anolis from Cuba. Proc. Biol. Soc. Washington 29: 19-20.

1919. Herpetology of Cuba. Mem. Mus. Comp. Zool. 47: 71-213.

BOULENGER, G. A.

1885. Catalogue of the lizards in the British Museum, (ed. 2), London
2: 1-497.
Cope, E. D.

1864. Contributions to the herpetology of tropical America. Proc.
Acad. Nat. Sci. Philadelphia 166-181.
Hardy, J. D.

1958. Tail prehension and related behavior in a New World lizard.
Herpetologica 14 : 205-206.

Ill IBAL, R.

1958. A preliminary investigation of the ecology and taxonomy of
Cuban lizards. Year Book Amer. Phil. Soc. 1957, 256-258.

1961. Thermal relations of five species of tropical lizards. Evolution
15: 98-111.
Ruibal, R. and E. E. Williams

1961. Two sympatric Cuban anoles of the carolinensis group. Bull.
Mus. Comp. Zool. 125: 181-208.
Smith, H. M.

1946. Handbook of lizards. Comstock Publishing Co. Ithaca, 1-557.

Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE
Vol. 125, No. 9

TAXONOMY OF THE DEEP SEA FISHES OF THE
GENUS CHAULIODUS

By James E. Morrow, Jr.

Department of Wildlife Management
University of Alaska, College, Alaska

CAMBEIDGE, MASS., U.S.A.
PEINTED FOE THE MUSEUM

October, 1961

Publications Issued by or in Connection
with THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 125.

Breviora (octavo) 1952 — No. 145 is current.

Memoirs (quarto) 1864-1D38 — Publication was terminated with

Vol. 55.

Johnsonia (quarto) 1941 -- A publication of the Department of
Mollusks. Vol. 4, no. 40 is current.

Occasional Papers of the Department op Mollusks (octavo)
1945 _ Vol. 2, no. 26 is current.

Proceedings op the New England Zoological Club (octavo)
1899-1948 — Published in connection with the Museum. Publication
terminated with Vol. 24.

Of the Peters "Check List of Birds of the World," volumes 1-3, 4
and 6 are out of print; volumes 5, 7 and 9 are sold by the Museum,
and future volumes will be published under Museum auspices.

Publications of the
Boston Society op Natural History

The remaining stock of the scientific periodicals of the Boston
Society of Natural History has been transferred to the Museum of
Comparative Zoology for distribution.

Proceedings — Volumes available: 3, 5, 6, 8, 11, 14-17, 20-22, 24-27,
30-34, 37. $4.00 per volume.

Occasional Papers : Volume 2, $5.00 ; Volume 3, $4.00 ; Volume 4
(1-3), $10.00; Volume 6, $5.00.

Memoirs : Requests for some specific memoirs can be filled but no list
is available.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 125, No. 9

TAXONOMY OF THE DEEP SEA FISHES OF THE
GENUS CHAULIODUS

By James E. Morrow, Jr.

Department of Wildlife Management
University of Alaska, College, Alaska

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

October, 1961

No. 9 — Taxonomy of the Deep Sea Fishes of the
Genus Chauliodus

By James E. Morrow, Jr.1

Representatives of the genus Chauliodus are found in all the
oceans of the world in temperate and tropical regions, and form
a rather closely knit group. Some species are widespread in their
distribution ; others appear to be confined to particular water
masses. Among the more widely ranging forms, populations ap-
pear to differ from one water mass to another, with the result
that a number of species and subspecies have been described.

The various forms of Chauliodus are not particularly well de-
fined in much of the literature, nor are they always easy to dis-
tinguish when specimens are at hand. The older descriptions, in
particular, made no allowance for normal variation within species,
and are, almost without exception, so brief and generalized that
it is well nigh impossible to determine what form was being des-
cribed. Yet the authors of the past cannot really be blamed for
this. Specimens were even more difficult to obtain in those days
than they are now, and taxonomic thought of the time was a far
cry from modern ideas.

In 1906, Brauer gave a summary, based on the literature, of
the species then known, but it was not until the expeditions of the
Dana that a series of specimens adequate for modern taxonomic
techniques became available. Basing their account on the speci-
mens collected in the Atlantic by the Dana expedition of 1920-22,
Regan and Trewavas (1929) produced the first modern work on
Chauliodus. They described a new species, C. danae, distinguish-
ing it from C. sloani chiefly on the basis of the more posterior
position of the dorsal fin and a different structure of the chin
barbel.

Later, Ege (1948) studied the large collections of Chauliodus
from the round-the-world Dana cruise of 1928-30. He described
two new subspecies of C. sloani (C. s. secundus from the Indo-
Pacific and C. s. schmidti from the eastern Atlantic) and reduced
several other species to the status of subspecies of C. sloani. Haff-
ner (1952b), studying the zoogeography of the genus, accepted
Ege's classification, though with some reservations as to the
propriety of the subspecific designations of some groups. Since
that time, virtually nothing has been done with the group as far

i Former address : Bingham Oceanographic Laboratory, Yale University.

250

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

as published information is concerned, although exploratory fish-
ing-, particularly in the north and central Pacific, has added a
great deal to our knowledge of the range of several species.

ACKNOWLEDGEMENTS

In carrying out the present work, we have had available to us
the large collections from the Dana Expedition 1928-30: the
Museum of Comparative Zoology, Harvard University ; the
United States National Museum; the Department of Ocean-
ography, University of Washington;1 and the Bingham Ocean-
ographic Collection, Yale University. In addition, we have been
able to examine other specimens from the British Museum
(Natural History); the Galathea Expeditions 1950-1952; the
Marine Laboratory, University of Miami ; the Scripps Institu-
tion of Oceanography; the Zoological Museum, Stanford Uni-
versity; the Department of Fisheries, University of Wash-
ington ; and the U.S. Fish and AVildlife Service Laboratory,
Honolulu. This material has been made available through the
kind cooperation of the following, to whom gratitude and thanks
are tendered : Dr. William Aron, Dr. E. Bertelsen, Dr. Henry B.
Bigelow, Dr. Anton Bruun, Mrs. M. Dick, Dr. Alfred Ebeling,
Dr. Robert Kanazawa, Dr. Ernest Lachner, Dr. Jorgen Nielsen,
Dr. C. R. Robins, Mr. R. Rosenblatt, Dr. W. F. Royce, Dr. F.
G. W. Smith, Dr. Leonard P. Schultz, Miss M. Storey, Dr. Ralph
Taylor, and Dr. Ethelwyn Trewavas.

DIAGNOSTIC CRITERIA

Early descriptions of species of Chauliodus depended in the
main upon the customary fin ray counts and body proportions.

Figure 1. Chauliodus sloani. Drawn from several specimens in the
Bingham Oceaaographic Collection by Shirley P. Hartman.

i The material from the University of Washington, Department of Oceanog-
raphy was collected by Dr. Wm. Aron. His work was supported by the National
Science Foundation and by the Office of Naval Research, Contract 477 (10).

MORROW: TAXONOMY OF CHAULIODUS 251

Garraan (1899) appears to have been the first to realize that the
number of serial photophores could be of some taxonomie impor-
tance in the group, in which he was followed by Brauer (1906).
Ege (1934), studying the related genus Stomias, discovered there
that the number and arrangement of the small body photophores
were characteristic for the various species. In his subsequent work
on Chauliodus (Ege, 1918), he continued with this same line of
thought, placing rather heavy emphasis on the number and type
of organ in each scale area, as well as utilizing other characters
such as the relative size of certain small body photophores, num-
ber and size of teeth, number of lower jaw denticles, etc. How-
ever, Ege appears to have based his conclusions with respect to
these more minute characteristics upon the examination of a
rather small number of specimens. Thus, in dealing with the
number and size of the small light organs in the various scale
areas, he describes only one or two specimens for each form.
Similarly, with respect to teeth and lower jaw denticles, he tabu-
lates data for only four or five specimens of each species. It
seemed necessary, therefore, to subject these and other matters to
a statistical examination, based on as many specimens as possible.

We therefore turn now to an examination of the various criteria
which have been used for the diagnosis of the several species of
Chaidiodus. For this examination, we have used as many speci-
mens as possible of each species. With respect to the serial photo-
phores, we have made use, wherever possible, of the data of other
authors as well as our own. It will be noted that there are dis-
crepancies here and there between the number of specimens listed
as study material and the number actually used in certain compu-
tations. This is due to the delicate nature of these fishes, with the
result that they are easily damaged in capture and preservation,
so that the character in question could not be observed with the
desired degree of accuracy.

Pre-anal distance less head length is measured from the
tip of the snout to the base of the first ray of the anal fin.
The head is measured from the tip of the snout to the pos-
terior edge of the opercular membrane, with the head as nearly
as possible in a normal position. Expressed as a percentage of
the standard length, this measurement has been used to set off
C. barbatus from all other species. However, as shown in Figure
2, barbatus is completely overlapped in this respect by macouni,
and to a great extent by sloani as well. To a lesser degree,
schmidti and pammelas also fall within range of barbatus for

252

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

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MORROW : TAXONOMY OF CHAULIODUS 253

this character. Although some of the differences which appear in
Figure 2 are certainly significant, both statistically and biologic-
ally, this character is really useful only to separate barbatus and
pammelas from danae.

Relative length of third and fourth premaxillary teeth has
also been used to separate barbatus from the other species. In
C. barbatus, and also in C. macouni, the third premaxillary
tooth is always longer than the fourth. In all the other species,
the opposite is true. Rarely in C. sloani and C. schmidti does
one find an individual whose premaxillary teeth do not fall
into the usual pattern, and the difference in the length of the
teeth is usually very slight in such instances. In nearly all the
species, there are also occasional individuals whose third and
fourth premaxillary teeth are of about equal length, but again,
these are rather rare (see Table I). The relative length of the
third and fourth premaxillary teeth, then, appears to be a fairly
good character for separating the two strictly Pacific Ocean forms
from the others.

Table I

Relative lengths of the third and fourth premaxillary teeth in
Chauliodus. The body of the table lists the number of
specimens of each species in each category.
Species

sloani

danae

schmidti

pammelas

barbatus

macouni

The number of light organs in each scale area of the second,
third and fourth rows has also been considered as diagnostic. In
the second row, each scale area generally has two organs in C.
sloani, C. pammelas, C. danae, and C. macouni. In C. schmidti
and some populations of C. sloani, there is a tendency for only a
single organ to be present, and in C. barbatus it is a rare indi-
vidual that has more than one much reduced organ present in
the second row areas. However, there is a great deal of variation,
particularly in C. sloani, where we have found 1, 2 or 3 organs in
these second row areas. Some indication of this degree of varia-
tion is shown in Table II, comparing C. sloani, C. schmidti and
the dannevigi population of C. sloani. It is apparent from this

3rd tooth

4th tooth

About

longer

equal

101

254 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

table that the number of small photophores in the second row is
not diagnostic for C. schmidti, although there is a tendency to-
wards reduction in this form. The same is true for the dannevigi
population of C. sloani. Particularly when it is realized that
many Mediterranean specimens match perfectly the descriptions
of dannevigi with respect to the arrangement of small photo-
phores, there seems no reason to consider dannevigi as anything
more than a population of C. sloani.

Table II

Distribution of the number of light organs in the scale areas
of the second row in three forms of Chauliodus. The body of
the table shows the number of specimens of each species with
the indicated percentage of second row organs containing two

small photophores.

Percentage of second row areas with two light organs

Species 0 10 20 30 40 50 (30 70 80 90 100

sloani 5443283557 21

schmidti 9611030010 0

dannevigi 2331005001 1

In the scale areas of the third row, there are always two small
photophores, except in C. barbatus. In that species there are three
or more, arranged in a cluster of one slightly larger organ with
two to many tiny organs. One specimen of C. oaroatus examined
by us appeared to have but two organs in about half of the third
row scale areas, but as the skin of this fish was in very poor condi-
tion, this appearance may have been due to damage. The number
of organs in the third row, then, will distinguish C. oaroatus from
all other species of the genus.

The scale areas of the fourth row each have two organs, as a
general rule, in all species except C. barb at us and C. macouni.
In these, the usual number is one. But in all forms, there is
a considerable amount of variation. In C. barbatus and C.
macouni, counts of fourth row organs vary from one to four, in
C. sloani from one to three. We regard the number of organs in
the fourth row scale areas as of only limited value in distinguish-
ing the species of Chauliodus.

The number of teeth in the lower jaw has several times been
used in the past in attempts to diagnose the species of Chauliodus,
but it requires only a glance at Table III to see that the number
of larger lower jaw teeth is of little taxonomic value. With such
a range of variation as is shown here, the best that can be done

MORROW: TAXONOMY OF CHAULIODUS 255

is to say that there are differences in the average number of teeth
in the lower jaw. C. sloani generally has six or seven teeth. C.
danae most frequently has seven or eight teeth in the lower jaw,
but there may be as few as five or as many as eleven. C. schmidti
most often shows six teeth, C. barbatiis and G. macouni six or
seven, and the few specimens of C. pammelas that have been ex-
amined or reported in the literature all had only five teeth in
the lower jaw.

Table III

Number of larger lower jaw teeth in various species of

Chauliodus. The body of the table shows the number of

specimens of each species. The data presented here include

also tooth counts from Ege (1948).

Number of Teeth

Species 5 6 7 8 9 10 11

sloani

danae

schmidti

barbatiis

macouni

pammelas

Also in the lower jaw, the number of small denticles that may
occur posteriorly, near the rictus, was used by Ege (1948) to
separate C. schmidti and C. pammelas. Both our own observa-
tions and Ege's published data indicate that in C. pammelas the
number of lower jaw denticles is a function of the size of the
specimen. With this limitation in mind, it can be seen from
Table IV that the number of lower jaw denticles can be a useful
secondary character to distinguish these two species from each
other, though it is of relatively little value amongst the others.

Table IV
in the lower jaw in Chauliodus*

3 4 .3 6 7 8 9 10

or
more
56871024
9 6 1

3 1 1
1 1
pammelas 1 1115

* Includes data published by Ege (1948) for 36 C. sloani, 7 C. danae, 4
C. barbatiis and 4 C. pammelas.

** Regan and Trewavas (1929) give 3 to S denticles for C. danae.

Number

denticles

Species

1 2

sloani

13 11

danae

3 4

schmidti

1 1

barbatiis

3 5

macouni

6 4

256 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

The relative lengths of the first and second premaxillary teeth
have heen indicated as a distinguishing feature by which C.
macouni might be separated from the other forms. The single
specimen of C. macouni taken by the Dana had a remarkably
short first premaxillary tooth, only 51.5 per cent of the length
of the second. In all other species, with a few rare individual ex-
ceptions, the first premaxillary tooth is 60 per cent or more of
the length of the second. This difference does not, unfortunately,
hold good. All of the specimens of C. macouni examined by us
have shown the first premaxillary tooth to be at least 60 per cent
of the length of the second. We have found short first teeth
only in C. barbatus. It may be of interest to note that all our
specimens of C. macouni came from the northern part of the
Pacific, whereas the Dana specimen was found in the tropics,
west of the Galapagos Islands. It may be that the short teeth
are somehow related to some characteristic of the Pacific Equator-
ial water mass, where C. barbatus also occurs. It may be men-
tioned here that the Dana specimen of C. macouni does not
appear to have been mis-identified. Dr. Ege kindly re-examined
the specimen, paying particular attention to the postocular
photophore. The elongate, pointed shape of this organ and its
location well behind the eye leave no doubt that it was correctly
named.

Ege (1948: 138, 139) erected a new subspecies of C. sloani —
C. sloani secundus — chiefly on the basis of slightly larger
light organs in the scale areas of the first and fifth rows in his
new form. He found that in C. sloani (sensu stricto) the small
organs of the first row had a diameter of 4.0 to 4.8 per cent of
the average length of the scale areas of the second row. In the
same species, the length of the largest organ in the scale areas
of the fifth row was 8.9 to 11.0 per cent of the average length of
the same scale areas in one group, 11.4 to 11.7 per cent in another
group. By contrast, the corresponding values for his new form
were given as 6.8 per cent for the dorsal row and 12.5 per cent
for the fifth row.

Examining specimens of C. sloani from the Atlantic, we ob-
served an extremely wide range of variation in these two charac-
ters, leading us to make a detailed study of the matter. The
length of the scale areas of the second row was measured on five
consecutive scales on each specimen, beginning at the fifth scale
behind the base of the ventral fins, and the average taken. The
diameter of the appropriate light organ of the corresponding
scales of the first and fifth rows was measured and expressed as

MORROW: TAXONOMY OF CHAULIODUS 257

a percentage of the average length of the scale areas of the sec-
ond row. This was done with 64 specimens of C. sloani from the
Atlantic, 12 specimens of C. s. secundus from the Indo-Pacific
area, and 13 specimens of C. sloani from either the same locali-
ties as the C. s. secundus or from nearby stations. There was
thus a total of 320 photophores from each row measured in the
Atlantic sample, and 60 or 65 in each of the others.

20 22 24 26

Figure 3. Distribution of the greatest diameter of the large light organ
in the fifth row, expressed as percentages of the average diameter of the
scale areas of the second row. A) C. sloani from the Atlantic. B) C.
sloani from the Indo-Pacific. C) C. sloani secundus.

Figure 3 shows the mean, range, ± two standard errors, and
± one standard deviation for the photophores of the fifth scale
row. There is obviously no significant difference in the value of
the mean for the three distributions. The distribution of C. s. sec-
undus is quite symmetrical, while the two samples of C. sloani
are skewed to the left. What significance, if any, this may have
we are not able to say.

2 4 6 8 10 12 14 16

Figure -4. Distribution of the diameter of the light organ in scale areas
of the first row, expressed as percentages of the average diameter of the
scale areas of the second row. Lettering as in Figure 3.

258

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

A similar plot for the photophore of the first row is shown in
Figure 4. Here there is an observable difference between the
mean value for C. s. secundus and the two samples of C. sloani.
However, the standard deviations of the distributions overlap
each other to a considerable degree, indicating a good deal less
than subspecific divergence.

The skewness of the Atlantic distribution, in particular,
arouses curiosity. The actual distribution of the light organs,
by size groups, has been plotted (Fig. 5), in order to obtain a
clearer comparison of the three samples. The two Indo-Pacific
samples look as though they were reasonably homogeneous, but
the plot of the Atlantic sample gives the impression that it may
be composed of two groups. One, the more numerous, has a mean
value somewhere around five; the other, with a greater range,
looks to have a mean in the neighborhood of nine. But we are
still unable to find any means of separating out two groups from
this single distribution. A scatter diagram of the size of the
organ of the first row plotted against the size of the organ of the

120

7 8

AVERAGE

9 10

DIAMETER

Figure 5. Frequency diagram of the distribution of the diameters of
the first row organs. Solid circles — C. sloani from the Atlantic. Open
circles — C. sloani from the Indo-Pacific. Triangles — C. sloani secundus.

fifth row (Fig. 6) shows in a rather striking fashion that there
is actually no real reason for attempting to make such a separa-
tion. There appears to be a general increase in the size of one
organ with the size of the other, but over so broad a base that it
is difficult to draw any hard and fast conclusions on this matter.

MORROW: TAXONOMY OF CHAULIODUS

259

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260 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Further, the measurements of the organs of Indo-Pacific C.
sloani and of C. sloani sccundus fall entirely within the scatter
of the Atlantic specimens of C. sloani. We are thus reinforced
in our conclusion that C. sloani sccundus should not be considered
as a subspecies.

The number of serial photophores in the lateral and ventral
series is an extremely valuable characteristic on which several
species may be distinguished. In particular, C. schmidti and C.
pammclas have much lower counts than any other species except
C. barbatus, but they are readily distinguished from C. barbatus
on other grounds. C. pammclas, with only 58 to 61 ventral organs
and 38 to 39 lateral ones, falls completely outside the range of
C. sloani for this character. For this reason we do not hesitate
to consider C. pammclas as a species completely distinct from C.
sloani. There seems to be no justification for maintaining C.
pammelas as a subspecies of C. sloani. The matter of C. schmidti
as a species or as a subspecies of C. sloani is not quite as clear
cut, for there is some degree of overlap between the highest
counts of C. schmidti and the lowest counts of C. sloani. Con-
fining the discussion of the Atlantic forms of C. sloani, we find
that this species has a total count of organs in the ventral row of
63 to 70. In the lateral row, the count is 42 to 48. For C. schmidti,
the corresponding values are 60 to 63 and 38 to 42. But we must
point out that in the ventral row counts, the number of C.
sloani that overlap the range of C. schmidti is 3 out of a total of
298 specimens. (We have here included data on 197 specimens
published by Ege (1948) and 101 specimens examined by us.)
For the 54 specimens of C. schmidti reported by Ege, 14 had 63
photophores in the ventral row, 40 had 60 to 62. The coefficient
of difference for the two distributions is 2.09, far above the
usual subspecific level and indicating a much greater degree of
differentiation. The same conclusion must be drawn from the
graphic presentation of the statistics of these data, shown in
Figure 7. The ranges of the two distributions meet at the value
63, but the two standard deviations are separated by a distance
equal to 1.6 times larger. We feel completely justified, then, in
raising C. schmidti to specific rank.

Two other items must be mentioned in this section. One of
these is the shape and location of the postocular photophore, a
characteristic which, as far as we know, has not previously been
utilized in this group. In all species except C. macouni, this
organ is more or less round and located generally somewhat an-
terior to a vertical through the posterior margin of the eye.

MORROW: TAXONOMY OF CHAULIODUS 261

In C. macouni, however, the organ is generally triangular and
pointed behind, sometimes notably elongate, and is placed pos-
terior to the vertical through the hind border of the eye.

C. SLOANI

SCHMIDTI

III! I I I I I I L

60 62 64 66 68 70

Figure 7. Distribution of the total number of serial photophores in the
ventral row in two species of Chaullodus.

The other characteristic is the structure of the barbel. This
was mentioned briefly by Regan and Trewavas (1929), but ap-
pears to have been ignored by most other workers. As Regan
and Trewavas pointed out, the barbel of C. sloani is tapered and
flexible, and disappears at a rather early age. This appears to be
true also of C. schmidti and C. pammelas. In C. danae and C.
barbatus, the barbel is stiff and compressed. In C. danae, the
organ is quite straight, without expansions, and has degenerated
in most specimens larger than 50 mm standard length. In C. bar-
batus, by contrast, the barbel bears a terminal, leaf -like expansion,
and the whole organ is retained even in the largest specimens. In
C. macouni the barbel resembles that of C. sloani, but is longer,
stiffened basally, and does not degenerate.

DEFINITIONS

Two terms that have been and will be used in this paper require
definition, as they are not ordinarily found in descriptions of
fishes.

Scale areas. These are actually the scale pockets. (For a
detailed description, see Morrow, Chauliodontidae, Fishes of the
Western North Atlantic, Vol. 3.) The scales of Chauliodus are
large and fairly heavy. However, they are extremely deciduous,
and appear also to dissolve readily in the ordinary preserving
fluids. In addition, the skin is, in life, covered with a thick layer
of mucus which makes it very difficult to distinguish the scales
themselves. Each scale pocket is outlined by a pigment pattern,

262 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

so that even after the scales are lost, the areas can often be dis-
tinguished with a fair degree of ease and accuracy. In the term-
inology used here, the most dorsal row of scales is called the first
row. The scales of this row are much smaller than the others,
and are often almost impossible to see. The most ventral row of
scales is termed the fifth row.

SM organs. These are the small light organs lying in the area
between the lateral and ventral rows of serial light organs. The
arrangement of these small organs is here indicated by a numer-
ical formula showing the number of small, unpigmented organs
at each end and the number of somewhat larger pigmented organs
in the middle of the series between adjacent pairs of large, serial
photophores. Thus, 1+3+0 would indicate one small unpigment-
ed organ at the anterior end of the series, three larger, pigmented
organs in the middle part, and no organ at the posterior end.

All descriptions of scale areas and SM organs refer to the part
of the body beginning four or five scales behind the bases of the
ventral fins and extending posteriorly for about ten scales.

Family CHAULIODONTIDAE

Characters. Body long, slender, compressed, covered by five
longitudinal rows of large scales. Scales deciduous, their loca-
tion often marked only by pigment pattern. Ventral fins before
middle of body measured from snout, of seven or eight rays.
Dorsal fin in anterior third of body, far in advance of anal fin,
its first ray much produced into a long filament, terminating in a
small flap. Adipose dorsal and anal fins present. Anal fin far
posterior, close to caudal.

Premaxillaries not protractile. Well developed epiotics pres-
ent beside supraoccipital. Parietals minute. First few vertebrae
without centra, the notochord enclosed in a sheath, first vertebra
with a large single parapophysis below, and long paired laminae
above representing neural arch. The enlargement of this vertebra,
together with the acentrous nature of the ones behind it, is re-
lated to throwing back the head in order to open the mouth in
feeding (Tchernavin, 1953). The remainder of the skeleton is
much like that of the Astronesthidae, and is but poorly ossified.

Snout short, more or less equal to eye. Nostrils large, immedi-
ately before eyes, the complex olfactory laminae exposed. Inter-
orbital less than eye, with a prominent bony ridge above each eye.
Teeth of premaxillary and mandible rigid, large to enormous,

MORROW: TAXONOMY OF CHAULIODUS 263

fang-like. Mandible with none to many tiny denticles near ric-
tus. Maxillary with numerous fine teeth on posterior half to two-
thirds of its ventral margin, entering posterior portion of gape.
Vomer toothless. Palatines with a few teeth anteriorly, followed
by a space, then several minute teeth. Operculum small. Four
gills, a slit behind the fourth. Gill arches armed with teeth, but
no true gill rakers. Mental barbel generally short and simple,
becoming much reduced or absent in juveniles and adults of
some species, retained in others.

Postocular luminous organ present, below and just before or
somewhat behind posterior margin of eye, another organ imbed-
ded in skin immediately before eye. Small photophores present
in each scale area. A row of large light organs present on each
side of body, beginning immediately behind gill opening and
ending above or nearly above anterior end of anal fin. Another
row of large photophores below these, beginning at anterior end
of isthmus and ending at caudal base. Between the two rows of
large organs is a wavy row of small ones (here termed SM or-
gans), whose pattern more or less repeats itself between each pair
of large serial organs, and appears to be diagnostic for some
species. Groups of small organs present on mid-ventral line be-
tween the two ventral rows of large photophores. Light organs
present on branchiostegal membranes, between rays.

Miscellaneous anatomy. The internal anatomy of the major
systems is typical of pelagic teleosts, with minor adaptations in
the digestive system which are presumably related to the availa-
bility of food in the mid-depths. It has been examined in detail
by Haffner (1952a), whence most of the following account has
been derived.

The most prominent feature of the digestive tract is the long,
sac-like stomach. Apparently it is not particularly distensible, but
achieves its capacity through its length. As with other pelagic
fishes, the esophagus, intestine and pyloric caeca join the stomach
at its anterior end.

The esophagus is muscular, lined with short columnar epithelial
cells, some of which, in the anterior portion, have a short, curved
spine at their free end. The spines disappear in the posterior
portion of the esophagus, and their function is unknown. In the
anterior portion of the stomach, the mucosal lining is thrown
into folds, with each fold bearing a lymph nodule at its tip. The
pyloric caeca are thin-walled, and the mucosa and submucosa
are much folded, resulting in the division of each caecum into

264 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

numerous small compartments. Haffner suggested that the com-
partments were so small as to prevent the entrance of food into
them, and hazarded that the caeca might be secretory in function.
The intestine, arising on the ventral side of the stomach at its
anterior end, extends straight back to the anus with no change in
diameter.

The kidneys are long and narrow, extending the full length of
the body cavity, and appear to be aglomerular. The urinary
bladder opens through a pore behind the opening of the genital
duct. The gonads resemble the kidneys in gross appearance, but
do not reach as far posteriorly. The genital ducts empty through
a pore between the anus and the urinary pore.

The swim bladder, according to Marshall (personal communi-
cation of information in press), is either absent, or degenerates
completely at a very early stage.

The microscopic structure of the photophores has been the
subject of several investigations, (Chiarini, 1900; Brauer, 1908;
Haffner, 1952a), with the result that they have been thoroughly
described and illustrated. There are four types, the simplest be-
ing merely a small, spherical mass of radially-arranged columnar
cells, without pigment layer, lens, or reflector. The next form, of
which the sub-ocular photophore is typical, also lacks lens and
reflector, but is provided with a layer of dark pigment covering
about % of its surface. Its structure resembles that of the sim-
plest type, except that instead of a lumen, the center of the organ
is filled with what Brauer interpreted as coils of cells, but which
Haffner thought could be explained more readily as the cut ends
of radially -arranged columnar cells. The more complex organs,
consisting of light-producing elements, pigment layer, lens and
reflector, are of two types, the bowl and cup-shaped organs, and
the bell-shaped organs. The former have a single, the latter a
double lens.

The photophores were early thought to be equipped with nerve
fibers, but later workers have interpreted the observed structures
as blood vessels, indicating that the luminescence of the photo-
phores is under hormonal rather than direct nervous control.

Range. The several species of Chauliodus, the only genus of
the family, are found in nearly all parts of the oceans, having
been taken throughout the North Atlantic and Mediterranean,
at various localities in the South Atlantic, and in the Pacific and
Indian oceans. In general, the range lies between 50°N and 40° S,
although C. macouni is commonly found in the Gulf of Alaska to

MORROW: TAXONOMY OP CHAULIODUS 265

60°N, and there are few records of C. sloani in the Atlantic be-
tween 55 °N and 65° N.

In their vertical distribution, the members of this group may
be found as close to the surface as 20 meters, and down to depths
as great as 2,800 meters. Several species appear to have different
depth preferences, and all appear to be more or less limited in
their distribution by certain physical and chemical characteris-
tics of the water masses, in general, larger individuals, and also
species reaching larger sizes, tend to live at greater depths than
do the smaller ones.

Genus CHAULIODUS Bloch and Schneider, 1801

Chauliodus Bloch and Schneider, Systema Ichthyologiae, Berlin,
1801: 430; type species C. sloani Bloch and Schneider, 1801, by
monotypy.
Generic Synonym :
Leptodes Swainson, Nat. Hist. Classification Fishes, Amphibi-
ans and Keptiles. London, 1839 : 298 ; type species L. sloanii
(Bloch and Schneider).
Generic characters. As for the family.

Size and habits. The various species included in Chauliodus
are of no more than moderate size. C. sloani has been recorded
at a length (standard or total?) of 350 mm (Zahl, 1953), but the
vast majority of individuals do not appear to exceed about 300
mm standard length. C. danae, in particular, does not appear to
reach more than about half this length.

The habits of this group, are, of course, unknown from direct
observation, but examination of stomachs of preserved individu-
als has shown them to be carnivores, feeding on other fishes and
on crustaceans. Tchernavin (1953) has deduced the detailed me-
chanics of their feeding actions from a study and dissection of
C. sloani. Larval development of C. sloani has been described, but
that of other species remains unknown. (For a resume, with
many references, of current knowledge of larval development in
Chauliodus, see Morrow, Fishes of the Western North Atlantic,
vol.3.)

Species. All told, some 14 names, which do not include com-
binations, have been applied to the members of the genus Chauli-
odus, but no one has ever attempted to accept all fourteen as
valid. The first attempt to list the species seems to have been that
of Garman (1899), who recognized five, but made no indication
of his thoughts on the remaining three which had been described

266 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

at that time. Brauer (1906) considered the matter in more detail
and accepted the same five species. He also pointed out that C.
setinotus Bloch and Schneider and C. schneicleri Risso were syn-
onyms of C. sloani Bloch and Schneider. Regan and Trewavas
(1929) gave detailed descriptions of their material of three
species and suggested that C. dannevigi and C. macouni might be
synonyms of C. sloani and C. barbatus, respectively. Finally, Ege
(1948) accepted only C. sloani, C. danae and C. barbatus as full
species, reducing pammelas, dannevigi and macouni to subspecies
of sloani and describing two new subspecies, schmidti and secun-
dum. Thus, he considered that there were three species, one of
which was split into six subspecies, for a total of eight forms.

In the present paper, we recognize six species. The subspecies
of Ege we feel should either be raised to specific rank or synony-
mized altogether, as has already been pointed out in the section
on species criteria.

Key to Species

la. Postocular photophore elongate, the exposed luminous
portion triangular or at least pointed behind ; the organ
itself located below or behind a vertical through posterior
edge of eye.

C. macouni Bean

Gulf of Alaska, North Pacific Ocean

lb. Postocular photophore round or nearly so, exposed lumin-
ous portion never triangular or pointed behind ; the organ
itself located below or before a vertical through posterior
edge of eye.

2a. Serial photophores of lateral series ventral to scale
areas of fourth row. Majority of scale areas of 3rd row
generally with a cluster of 3 or more small light organs.
C. barbatus Garman
Gulf of Panama, tropical Pacific

2b. Serial photophores of lateral series below scale areas
of fifth row. Scale areas of 3rd row with not more than
2 small light organs.

3a. Dorsal origin over 9th to 12th OV photophore.
4a. Posterior unpigmented organ of each SM
series generally absent, or at least much re-
duced. SM formula generally 1+2+0. Chin
barbel (when present) short, stiff, compressed,

MORROW: TAXONOMY OF CHAULIODUS 267

absent in most specimens larger than ca 50 mm.
C. danae Regan and Trewavas
North and South Atlantic Ocean.
4b. Posterior organ of each SM series nearly as

large as the central organs. SM formula

generally 1+3+1 or 1+4+0. Barbel tapered,

slender, flexible, generally present in most

specimens up to ca 100 mm.

C. sloani Bloch and Schneider
(some Indo-Pacific variants)
3b. Dorsal origin over 5th to 8th OV photophore.
5a. Total ventral photophores 64 to 72, rarely

62 or 63. Total lateral photophores 43 to 48,

rarely 42.

C. sloani Bloch and Schneider
Atlantic, Pacific, Indian oceans,
Mediterranean Sea.

5b. Total ventral photophores 58 to 63. Total

lateral photophores 38 to 42.

6a. SM series with 3, sometimes 4 larger

pigmented organs in each series. Total

ventral organs 58 to 61, total lateral organs

38 to 39.

C. pammelas Alcock
Northern Indian Ocean
6b. SM organs with only 2 larger pigmented
organs in each series. Total ventral or-
gans 60 to 63, total lateral organs 38 to 42.
C. schmidti Ege
Eastern Atlantic Ocean

Chauliodus barbatus Garman, 1899

Study material. Nineteen specimens, 74 to 183 mm standard
length, from the Gulf of Panama, Galapagos Islands and off Peru.

Distinctive characters. C. barbatus is particularly character-
ised by the following: Serial photophores of the lateral series
located on verticals passing approximately through the centers of
the scale areas of the fourth row; barbel stiff, compressed, ex-
panded at tip, present at all stages ; dorsal origin over 8th to 11th
OV photophore; total lateral photophores 38-41; total ventral
photophores 59-63.

268 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Description. Proportional measurements of the study material
given as average percentages of standard length except as indi-
cated otherwise, with the range of variation shown in parentheses.

Body: depth 9.9 (7.1-14.8).

Head: 15.3 (13.5-16.5).

Eye: 3.1 (2.6-3.4); 20.17o of head (17.0%-22.3% of head).

Snout: 3.5 (3.1-4.1) ; 22.6% of head (19.5%-26.0% of head).

Distance from snout : to origin of dorsal fin 27.0 (25.1-29.0) ; to
origin of anal fin 81.7 (79.0-83.1) ; to origin of ventral fin
41.9 (40.5-44.1).

Pre-anal length without head : 66.5 (63-69) .

Dorsal fin : rays 6.

Anal fin : rays 10-13.

Pectoral fin : rays 10-13.

Ventral fin: rays 7.

Vertebrae : 51-55, most often 53.

Serial photophores : Ventral row : IP 9-11 ; PV 16-19 ; VAV 22-
24; AC 10-11; Total 59-63. Lateral row: OV 17-18; YAL 20-23;
Total 38-41.

Body elongate, slender, compressed, the depth averaging about
10 per cent of the standard length, slightly deeper than in other
species. Barbel present at all stages, short, stiff, compressed,
terminally expanded. Edges of expanded part usually crenate
(Pig. 8).

Head about % of standard length, with prominent bony ridges
above each eye. Eye round, about % of head. Snout slightly
longer than eye diameter. Subocular organ present below anter-
ior part of eye, deeply embedded in skin. Postocular organ more
or less round, located below posterior part of eye.

Mouth large, jaws almost equal to head length. Premaxillaries
with four teeth, second tooth longest, third tooth longer than
(rarely equal to) fourth. Many small oblique teeth on maxillary.
Mandible with 6 to 9 larger teeth, and up to 5 tiny denticles pos-
teriorly, near corner of mouth.

Origins of pectoral fins low on body, just anterior to posterior
edge of opercular flap when head is in normal position. Fins of
10 to 13 rays. Ventral fins of 7 rays, their origins before middle
of standard length. Dorsal fin of 6 rays, its origin over the 8th
to 11th OV photophore. Anal fin far behind dorsal, close to
caudal, with 10 to 13 rays, most often 12 or 13. Caudal fin forked.

Scale areas of the second row almost always, and of the fourth
row usually, with only one small light organ. Areas of the third

MORROW : TAXONOMY OF CHAULIODUS

269

row with a cluster of three or more small organs, usually one
slightly larger and two to many rather minute ones. Serial photo-
phores lying below scales of fourth row, scales of fifth row extend-
ing ventrally between the organs of the lateral row. SM organs
small, usually five to seven of about equal size in each section.

5 MM

I MM

Figure 8. Barbel of C. barbatus Garman, drawn from Galathea specimen
No. 15. A) Showing relation to lower jaw. B) The barbel itself, much
enlarged.

Size. The largest specimen examined was 183 mm in standard
length. It seems likely that this species does not much exceed
200 mm.

Relationships. C. barbatus clearly represents an individual
offshoot within the genus. Although the stiff compressed barbel
suggests affinities with C. danae, and the retention of the barbel
throughout life is found also in C. macouni, the arrangement of
the serial light organs below the fourth, rather than the fifth,

270 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

scale row, the presence of a group of organs rather than a pair in
the areas of the third row, and the increased number and reduced
size of the SM organs all indicate that C. barbatus is the most
divergent species of the genus.

Range. Known only from the eastern part of the Pacific Equa-
torial water mass, from the coast of Peru north to the Gulf of
Panama (possibly as far north as Central America) and west-
ward to about 100°W. In depth, C. barbatus has been taken down
to about 1200 fathoms, but the majority of records seem to be
from between 500 and 700 fathoms.
Synonyms and references:

Chauliodus barbatus Garman, Mem. Mus. Comp. Zool. Harvard,
24, 1899: 271-273, PI. K, figs. 2, 2a (type descr., illus., type
localities 6°22'20"N, 81°52'W, 465 fath., and 3°09'N, 82°08'W,
1132 fath., type specimens, Harvard Mus. Comp. Zool., Nos.
28489 and 28490) ; Brauer, Ergebn. Deutsch. Tiefsee-Exped.
"Valdivia", 15, Syst. Teil, 1906: 38-40 (comparison species) ;
Gilbert, Proc. U.S. Nat. Mus., 48, 1915: 321 (name) ; McCul-
loch, Biol. Res. "Endeavour", 1916, 4 (4): 181 (name);
Regan and Trewavas, Dana Rept., No. 5, 1929: 38-39 (descr.,
Gulf of Panama) ; Jordan, Evermann and Clark, Rep. U.S.
Comm. Fish., (1928) 1930, App. X: 71 (name); Ege, Dana
Rept., No. 31, 1948: 148 pp. (descr., synonymy, zoogeogr.) ;
Haffner, Syst. Zool., 1 (3), 1952: 114 (name, Gulf of Panama
to Galapagos) ; Marshall, Aspects of Deep Sea Biology, New
York, 1954: 65 (name).

Chauliodus danae Regan and Trewavas, 1929

Study material. Seventy specimens 37 to 133 mm standard
length, from the Caribbean Sea and North Atlantic Ocean.

Distinctive characters. The posterior position of the dorsal
origin, the short stiff barbel of the young, and the reduced SM
organs serve to distinguish C. danae.

Description. Proportional measurements of the study material
expressed as mean percentages of standard length except as
noted otherwise, with the range of variation shown in paren-
theses. Data from other authors in brackets.

Body: depth 7.7 (4.7-11.9).

Head: 12.9 (11.9-14.3).

Eye: 3.3 (2.2-4.9); 26.970 of head (17.7%-43.2% of head).

Snout: 2.8 (1.7-3.7) ; 22.3% of head (16.7% -28.8% of head).

MORROW : TAXONOMY OF CHAULIODUS

271

Distance from snout: to origin of dorsal fin 28.1 (25.9-32.3) ;
to origin of anal fin bo. 7 (83.5-88.7) ; to origin of ventral
fin 12.7 (39.7-45.7).
Prc-anal length without head: 73 (71-77).
Dorsal fin: rays 6.
Anal fin: rays 10-12.
Pectoral fin: rays 12-13 [14].
Ventral fin: rays 7.
Vertebrae: 51-57, most often 53-55.
Serial photophores: Ventral row:
VAV[22]23-26; AC 8-10; Total
17-20; VAL 22-25; Total 40-44.
Body compressed, elongate, slender, the depth averaging only
about 1/15 of the standard length. Barbel present only in young
(less than ca 50 to 55 mm SL), compressed, straight, with a
stiffening axial rod, reduced to a small triangular stump in
adults. (Fig. 9.)

IP 9-10; PV[ 17] 18-21;
61-65. Lateral row: OV

I MM

.,'-- "> ..■..JL i ,,.(.. ......

3MM

Figure 9. Barbels of C. danac. A) Lateral view, with part of lower
jaw cut away, of barbel of a 50 mm specimen (Bingham Oceanogr. Coll.,
No. 2938). B) Ventral view of same. C) Ventral view of barbel of a
114 mm specimen (Bingham Oceanogr. Coll., No. 2935) showing reduction
in adult.

Head averaging about 3/s °f standard length, bony ridges
present above eyes. Eye round, its diameter quite variable,
generally somewhat larger in males than in females (Regan

272 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

and Trewavas, 1929: 34). Snout slightly shorter than eye
diameter. Suboeular organ present below front of eye, deeply
embedded. Postocular organ round, below posterior part of eye.

Mouth large, jaws almost equal to head length. Premaxillaries
with four teeth, second longest, third tooth shorter than (rarely
about equal to) fourth. Many small, oblique teeth on maxillary.
Mandible with 5 to 11 teeth, generally 7 or 8, with up to 8
denticles posteriorly near corner of mouth (see Table IV).

Pectoral fins with 12 to 14 rays, their origins low on body,
about below posterior edge of opercular flap. Ventral fins with
7 rays, their origins before middle of standard length. Dorsal
fin of 6 rays, its origin generally over 9th to 11th OV photo-
phore, rarely 8th or 12th. Anal fin far behind dorsal, close to
caudal, with 10 to 12 rays. Caudal fin forked.

Scale areas generally clearly marked by pigment patterns,
each area with one or more small light organs, those of the third
row with not more than two photophores. Areas of second and
fourth rows each with two organs. Serial photophores lying
below scales of fifth row. SM organs generally with the posterior
organ of each section absent or at least much reduced, the SM
formula 1+2+0.

Size. The largest specimen seen by us was 133 mm in standard
length. According to Regan and Trewavas (1929: 35), the
maximum size is about 140 mm.

Relationships. Closest to C. sloani, from which it differs chiefly
in the structure of the barbel, the posterior placement of the
dorsal fin, the reduced SM organs and the somewhat lower
number of serial photophores.

Range. The majority of the known specimens of C. danae
have come from a broad east -west belt across the North Atlantic,
extending roughly from 20°N to 40°N. However, other speci-
mens have been recorded from the South Atlantic, as far as
33°53'45"S, and from as far north as 51°N. In addition, the
species has also been recorded from the area along the eastern
edge of the Caribbean Sea and from the Yucatan Channel, west
of Cuba. Within these areas, it appears that C. danae is re-
stricted to water in which the degree of oxygen saturation is
at least 50 per cent. Depth wise, C. danae has been recorded as
far down as about 3500 meters (7000 meters of wire out), but
the majority of specimens appear to be taken in the upper 500
meters of water.

MORROW : TAXONOMY OF CHAULIODUS 273

Synonyms and References :

Chauliodus danae Regan and Trewavas, Dana Rept., No. 5, 1929 ;
34-38, pi. YII (type descr., type locality 13°03'N, 59°50'W,
300 m wire, type specimen Dana Coll. No. St. 1182) ; Norman,
Discovery Rept., 2, 1930: 308 (South Atlantic); Borodin,
Bull. Mus. Comp. Zool. Harvard, 72 (3), 1931: 64 (Bermuda) ;
Fowler, Bull. Amer. Mus. Nat. Hist., 70 (2), 1936: 1199-1200
(name); Beebe, Zoologiea, N.Y., 22 (14), 1937: 201 (Ber-
muda, 300-1,000 fath.) ; Parr, Bull. Bingham Oceanogr. Coll.,
3 (7), 1937 ; 58 (Bahamas, Bermuda) ; Bertin, Bull. Mus. Hist.
Nat. Paris, (2) 11, 1939: 382 (name); Nybelin, Goteborg
Vetensk. Samh. Handl., (B) 5 (16), 1948: 30 (N.E. Atlantic) ;
Ege, Dana Rept., No. 31, 1948: 1-148 (classification, phy-
logeny, zoogeography) ; Haffner, Systematic Zool., 1, (3),
1952: 112-113 (zoogeography); Grey, Fieldiana:Zool., 37,
1955: 277-278 (Bermuda, stomach contents, eggs); Koefoed,
Rep. Sars N. Atlantic Deep Sea Exped., 4 (2) (5), 1956:
19-20 (N. Atlantic, table of measurements).

Chauliodus atlantis (partim) Barbour, Proc. New Engl. Zool.
CI., 19, 1942: 46.

Probable synonym :

Chauliodus pammelas Pappenheim, Deutsch. Sudpolar Exped.,
15, Zool. 7, 1914: 167 (N. Atlantic).

Not Chauliodus danae Phillipps, Rec. Dominion Mus., 1 (1),
1942: 53-54 (Cook Strait).

Chauliodus macouni Bean, 1891

Study material. Ninety-three specimens, 30 to 182 mm stand-
ard length, from the northeastern Pacific Ocean and south of
Japan.

Distinctive characters. The generally triangular form of the
postocular organ, and iti location behind and below the eye
provide absolutely distinctive characters by which to distinguish
C. macouni from all other species of the genus.

Description. Proportional measurements of the study material
expressed as average percentages of standard length, unless indi-
cated otherwise, with the range of variation shown in paren-
theses. Data in brackets from other authors.

Body: depth 8.1 (6.7-10.1).

Head: 15.8 (13.2-17.2).

Eye: 3.3 (2.8-4.0) ; 20.87c of head (16.9%-28.1% of head)
[30.3].

274

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Snout: 3.8 (2.8-4.5) ; 24.4% of head (17.7%-28.7% of head)

[30.3].
Distance from snout: to origin of dorsal fin 24.9 (22.2-27.8) ;
to origin of anal fin 83.3 (80.9-84.7) [76.8-90.8] ; to origin
of ventral fin 41.3 (37.6-44.5) [35.7-47.5].
Pre-anal length without head: 68 (64-72).
Dorsal fin: rays 6 [rarely 7].
Anal fin: rays 10-13.
Pectoral fin: rays 10-13.
Ventral fin: rays 7 [rarely 8].
Vertebrae: 56-62.
Branchiostegal rays: 16-20 [21].

Serial photophores: Ventral row: IP 9-12; PV 17-21; VAV

26-29 [30] ; AC 10-13; Total 66-69. Lateral row: OV 17-20;

VAL 24-28 [29] ; Total 43-46.

Body slender, elongate, compressed, its depth averaging about

1/12 of the standard length. Barbel rather longer than in other

species, tapered, stiff basally but flexible near tip, present at all

stages (Fig. 10).

5MM

Figure 10. Barbel of C. macouni. Drawn from Bingham Oceanogr. Coll.,
No. 1287, 159 mm standard length.

Head almost 1/6 of standard length, with the usual bony
ridges above eyes. Eye round, its horizontal diameter averaging
about 1/5 of head. Snout generally a little longer than eye.
Subocular organ present below and before eye, deeply em-
bedded. Postocular organ located generally behind a vertical
through posterior margin of eye, the organ itself distinctly

MORROW: TAXONOMY OF CHAULIODUS 275

elongate, the exposed luminous surface generally markedly tri-
angular or at least pointed behind, sometimes notably attenuate.

Mouth large, jaws almost equal to head. Premaxillaries with
four teeth, second largest, third tooth longer than (rarely
equal to) fourth. Many small, oblique teeth on posterior part
of maxillary, which enters gape. Mandible with five to nine
larger teeth, generally six or seven. Usually no denticles on
posterior part of mandible, but sometimes as many as four
present. (See Table IV.)

Pectoral fins arising low on body, below posterior edge of
opercular flap, of 10 to 13 rays. Ventral fins of 7 rays, arising
well before middle of standard length. Dorsal origin over 6th
to 9th OV photophore, average predorsal distance about *4 of
standard length. Anal fin far behind dorsal, close to caudal, of
10 to 13 rays. Caudal fin forked.

Body with five longitudinal rows of scales, large serial photo-
phores of lateral row lying below scales of fifth row. Scale
areas of second row generally with only a single light organ,
those of third row with two, those of fourth row usually with
one photophore. SM organs generally 0+2+1, but showing
considerable variation, sometimes 1+1+0, the latter formula
found in the type specimen and a few others.

Size. The largest specimen examined, at 182 mm SL, appears
to be the longest on record, suggesting that this species may
not ever grow to much over about 200 mm SL.

Relationships. C. macouni appears to be closer to C. sloani
than to any other species in the genus, although the long third
tooth of the premaxillary and the permanent nature of the
chin barbel indicate affinities with the stock from which C.
barbatus must have arisen.

Range. The vast majority of records of C. macouni have come
from the subarctic Pacific water mass and the intermediate
water to the west of it, with a few from the transitional region
that extends south along the California coast. There are a few
records from other masses, e.g., one from the Pacific Equatorial
Water near the Galapagos Islands and another from south of
Japan in the border region between Intermediate and "Western
North Pacific Central Water, but it is quite possible that these
were strays and that these records do not really represent the
normal extension of the range. Present information, then, sug-
gests that the ordinary range of C. macouni includes the south-
ern part of the Bering Sea, southeastward through the Gulf of

276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Alaska at least as far as the southern part of California, and

westward about to Sakhalin, possibly as far as Japan.

Synonyms and references :

Chauliodus macouni Bean, Proc. U.S. Nat, Mus., 13, (1890)
1891: 44 (type descr., type locality 51°23'N, 130°34'W, 876
fath., type specimen U.S. Nat. Mus., No. 45372) ; Gilbert,
Rept. U.S. Comm. Fish., (1893) 1895: 402 (off Oregon,
Alaska) ; Goode and Bean, Oceanic Ichthyol., Washington,
1895: 513 (name); Jordan and Evermann, Bull. U.S. Nat,
Mus., 47, 1896: 585 (descr.); Garman, Mem. Mus. Comp.
Zool. Harvard, 24, 1899: 274 (name, meristic counts) ; Brauer,
Ergebn. Deutsche Tiefsee-Exped. "Valdivia", 15 Syst. Teil.,
1906: 38-40 (comparison other spp.) ; Gilbert, Proc. U.S. Nat.
Mus., 48, 1915: 321-322 (descr., comparison other spp., So.
Calif.); McCulloch, Biol. Res. "Endeavour", 4 (4), 1916:
181 (name) ; Jordan, Evermann and Clark, Rep. U.S. Comm.
Fish., (1928) 1930, App. X: 71 (name) ; Chapman, Occ. Pap.
B. C. Prov. Mus., No. 2, 1940: 5-11 (distrib., descr., compari-
son C. barbatus) ; Clemens and Wilby, Bull. Fish. Res. Bd.
Canada, 68, 1946: 108-109 (descr., illus., food, range) ; Barra-
clough, Copeia, 1950 (3) : 241-242 (inshore record, Brit.
Columbia) ; Copeia, 1954 (1) : 75-76 (same).

Chauliodus cmmclas Jordan and Starks, Bull. U.S. Fish. Comm.,
22, 1904: 579 (type descr., type locality Sagami Bay, Japan,
120-265 fath., type specimen U.S. Nat, Mus., No.' 51464) ;
McCulloch, Biol. Res. "Endeavour", 4 (4), 1916: 181 (name).

Chauliodus sloanei macouni Ege, Dana Rept., No. 31, 1948:
148 pp. (descr., near Galapagos Is.) ; Haffner, Systematic
Zool., 1 (3), 1952: 114, 132 (name, range).

Chauliodus pammelas Alcock, 1892

Study material. Seven specimens, 25 to 159 mm standard
length, from the Gulf of Aden, Arabian Sea and near the
Maldive Islands.

Distinctive characters. C. pammelas is set off primarily by
the low number of serial photophores, the forward position of
the dorsal fin, and the 3 or 4 pigmented SM organs.

Description. Proportional measurements of the study material
as mean percentages of standard length, unless indicated other-
wise, with range of variation in parentheses.

Body: depth 8.5 (7.2-9.5).

MORROW : TAXONOMY OF CHAULIODUS 277

Head: 15.1 (14.0-16.5).

Eye: 3.3 (2.8-4.4) ; 21.6% of head (19.1%-23.4% of head).

Snout: 3.5 (3.1-4.4) ; 22.3% of head (20.6%-25.1% of head).

Distance from snout: to origin of dorsal fin 23.2 (21.5-26.7) ;
to origin of anal fin 83.5 (82.5-84.5) ; to origin of ventral
fin 43.8 (41.5-45.3).

Pre-anal length without head: 67.9 (63-70).

Dorsal fin: rays 6.

Anal fin: rays 12-13.

Ventral fin: rays 7.

Pectoral fin : rays 12-14.

Vertebrae: 50-53.

Branchiostcgal rays: 17-20.

Serial photophores: Ventral row: IP 10; PV 17-19; VAV
21-23; AC 9-11; Total 58-61. Lateral row: OV 18-19; VAL
19-22. Total 38-39.

Body, slender, elongate, compressed, depth about 1/12 of
standard length. Chin barbel reduced to triangular stump in
study material.

Head 1/6 to 1/7 of standard length, a bony ridge above each
eye. Eye round, average diameter a little more than 1/5 of
head, averaging slightly less than snout length. Snout short, less
than !/4 of head. Subocular organ present below front of eye,
deeply embedded in skin. Postocular organ round, below eye,
before a vertical through posterior margin of eye.

Mouth large, jaws nearly equal to head length. Premaxil-
laries with four teeth, second tooth longest, fourth tooth longer
than third. Maxillaries with numerous small, oblique denticles
on posterior part of ventral margin, entering gape. Mandible
with five teeth, and up to 17, perhaps more, tiny denticles near
corner of mouth. The number of denticles appears to be a
function of the size of the individual, increasing as the fish
grows.

Origins of pectoral fins low on body, about below posterior
edge of opercular flap. Fins of 12 to 14 rays. Ventral origins
before middle of standard length, close together at mid-ventral
line. Dorsal fin arising over 6th or 7th lateral OV photophore,
pre-dorsal distance 21.5 to 26.7 per cent of standard length.
Anal fin far behind dorsal, close to caudal, with 12 or 13 rays.
Caudal fin forked.

Body with five longitudinal rows of scales, the scale areas
marked off by pigment patterns. Second row with one or (more

278 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

often) two small organs in each scale area. Each area of third
row with only two small light organs, areas of fourth row also
with two organs. Serial photophores ventral to scale areas of fifth
row. SM organs moderate, SM formula usually 0+3+1, some-
times 0+4+0.

Size. The type specimen, described by Alcock (1892: 355)
as "10 inches long," is the largest of which we have found a
record.

Relationships. C. pammelas is obviously derived from the
sloani stock, and appears to occupy a position close to C. schmidti
of the Atlantic, which it resembles, especially in the low number
of serial photophores. Whether the ecological positions are
similar is not known.

Range. This species has been recorded from the Gulf of
Aden, eastwards across the Arabian Sea to the Maldive-Lacca-
dive area, south and west of Ceylon, in the Bay of Bengal and
southwest of the Nicobar Islands, and once from the Flores
Sea. The last is the only record ascribed to this species south
of the equator. The depth range apparently goes to something
on the order of 2500 meters, with several other positive records
between 500 and 1000 meters, approximately. Unfortunately,
this species is not particularly abundant in collections, and
its distribution is therefore not well known.
Synonyms and references:

Chauliodus pammelas Alcock, Ann. Mag. Nat. Hist., (6) 10,
1892: 355 (type descr., type locality Laccadive Sea, 8°49'N,
73°18'45"E, 1370 fath., type specimen Indian Mus. No. 13183) ;
Goode and Bean, Oceanic Ichthyol., Washington, 1895: 96
(name) ; Alcock, Descr. Cat. Indian Deep Sea Fishes, Cal-
cutta, 1899: 145 (not seen) ; Illus. Zool. "Investigator," 1900:
PI. 30, fig. 4 (not seen) ; Brauer, Ergebn. Deutsche Tiefsee-
Exped. "Valdivia", 15 Syst. Teil, 1906: 38-42 (comparison
other spp.) ; 42-43 (descr.); Weber and de Beaufort, Fishes
Indo- Austral. Archipel., Leiden, vol. 2, 1913 : 110-111 (name) ;
McCulloch, Biol. Res. "Endeavour", 4 (4), 1916: 181
(name); Regan and Trewavas, Dana Rept., No. 5, 1929: 31
(name) ; Fowler, Bull. Amer. Mus. Nat, Hist,, 70 (1), 1936:
221 (name) ; Norman, Sci. Rept. John Murray Exped., 7(1),
1939: 21-22 (descr., distrib.).
Chauliodus sloanci pammelas Ege, Dana Rept., No. 31, 1948:
148 pp., (descr., synonymy, zoogeogr.) ; Haffner, Systematic
Zool., 1(1), 1952: 113-133 (zoogeogr.).

MORROW: TAXONOMY OF CHAULIODUS 279

Chauliodus sloani pammelas, Marshall, Aspects Deep Sea Biol.,

N.Y., 1954: 66 (name).
Probable synonym :
Chauliodus sloanii (partim), Alcock, Descr. Cat. Indian Deep

Sea Fishes, Calcutta, 1899: 144 (not seen).
Not Chauliodus pammelas Pappenheim, Deutsche Siidpolar Ex-

ped., 15, Zool. 7, 1914: 167 (name, N. Atlantic).

Chauliodus sohmidti Ege, 1948

Study material. Thirty-five specimens, 49 to 199 mm standard
length from the eastern Atlantic.

Distinctive characters. Chauliodus schmidti is distinguished
by the forward position of its dorsal tin, the low number of
serial photophores, and the presence of only two larger pig-
mented light organs in each section of the SM series.

Description. Proportional measurements of the study material
expressed as mean percentages of standard length unless noted
otherwise, with the range of variation in parentheses.

Body: depth 8.4 (6.3-10.7).

Head: 13.8 (12.2-14.9).

Eye: 3.1 (2.4-4.1); 22.4% of head (18.4%-27.8'/(' of head).

Snout: 3.3 (2.8-3.7) ; 23.9% of head (20.4%-26.6% of head).

Distance from snout: to origin of dorsal fin 21.4 (19.4-23.4) ;
to origin of anal fin 84.5 (82.8-85.8) ; to origin of ventral
fin 42.9 (39.9-46.0).

Pre-anal length without head: 71 (68-72).

Dorsal fin: rays 6, rarely 5.

Anal fin: rays 10-12.

Pectoral fin: rays 11-13.

Ventral fin: rays 7.

Vertebrae: 52-55 [56].

Branchiostegal rays: 17-19.

Serial photophores: Ventral row: IP 9-11; PV 17-20; VAV
22-25; AC 9-11; Total 60-63. Lateral row: OV 17-20; VAL
21-24; Total 38-42.

Body elongate, slender, compressed, the depth averaging about
1/12 of the standard length. Chin barbel reduced to a stump, at
least in adults.

Head 1/8 to 1/7 of standard length, bony ridges present
above eyes. Eye round, its diameter averaging a little more
than 15 head length. Snout generally slightly longer than eye.

280 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Subocular organ present below front of eye, deeply embedded.
Postocular organ round, below posterior part of eye.

Mouth large, jaws almost equal to head. Premaxillaries with
four teeth, second tooth longest, fourth tooth generally longer
than third. Maxillaries with numerous oblique denticles on pos-
terior part of ventral margin, entering gape. Mandibles with
five to nine (most often six) larger teeth, and none to two small
denticles posteriorly near corner of mouth.

Pectoral fins arising low on body, below posterior part of
opercular apparatus, of 11 to 13 rays. Origins of ventral fins
close together at midventral line, before middle of standard
length, fins with 7 rays. Dorsal with six, rarely five rays, dorsal
origin over 5th to 7th OV photophore. Anal origin far behind
dorsal, close to caudal, fin with 10 to 12 rays. Caudal fin
forked.

Sides of body with five longitudinal rows of scales, the scale
areas marked by pigment patterns. Each scale area with one
to several small photophores, those of the second row generally
with one, sometimes two, of the third and fourth rows with two
such organs. Serial photophores located ventral to scale areas
of fifth row. SM organs with two larger, pigmented organs in
each section, SM formula 1+2+1.

Size. The largest known specimen is something over 200 mm
long.

Relationships. C. schmidti is very close to C. sloani of the
Atlantic, differing from that group in the lower number of serial
photophores, and in a tendency for the scale areas of the second
row to have only a single small light organ more often than
two (see Table II). It would seem that C. schmidti has probably
developed directly from the Atlantic population of C. sloani.

Range. C. schmidti apparently occurs only in the eastern
Atlantic, off the coast of Africa, between approximately 10°S
and 20°N. Its east-west distribution is as yet unknown. The
species has been found at depths as shallow as 25 meters and as
great as 3000 meters. The Dana records (Ege, 1948) show that
of their 96 specimens, approximately 48 per cent were taken
at depths of 50 meters or less, and about 40 per cent came from
between 500 and 1500 meters. There seems to be a strong
tendency for small individuals, 50 mm or less, to occur above
the 500 meter depth, while specimens over 75 mm are most
frequently found at depths of 500 meters and more.

MORROW: TAXONOMY OF CHAULIODUS 281

Synonyms and references :

Chauliodus sloanei schmidti Ege, Dana Rept., No. 31, 1948: 139
(type closer., type locality "Tropical waters off W. coast of
N. Africa; north-eastern South Atlantic, to ca. 8°S."; type
specimen Dana Coll. No. St. 1005) ; Haffner, Systematic Zool.,
1(1), 1952: 113-133 (zoogeogr.).

Chauliodus sloani schmidti Marshall, Aspects of Deep Sea Biol-
ogy, N.Y., 1954: 66 (name).

Probable synonym :

Chauliodus sloanei (partim) Norman, Discovery Rept., 2, 1930:
308.

Chauliodus sloani Bloch and Schneider, 1801

Study material. Three hundred and sixty specimens, 25 to 278
mm in standard length, from the Atlantic Ocean, Gulf of Mexico,
Mediterranean Sea, eastern, middle and western Pacific Ocean,
and the Indian Ocean.

Distinctive characters. Because of the rather wide variation
between populations of C. sloani, it is difficult to give a succinct
summary of its distinctive characters. In general, C. sloani may
be characterized by a greater number of serial photophores than
any species except C. maeouni, a round postocular organ, and
the origin of the dorsal fin over the 5th to 8th OV photophore
except in some variants from the Indo-Pacific region. In these,
however, if the dorsal origin is over the 9th or 10th OV photo-
phore, the SM organs are 1+3+1 or 1+4+0 in each section,
which will separate them from C. danae of the Atlantic.

Description. Proportional measurements of the study material
given as mean percentages of standard length, except as noted,
with the range of variation in parentheses. Data in brackets
from other authors.

Body: depth 7.9 (5.6-12.0).

Head: 13.6 (10.5-16.3).

Eye: 3.1 (2.0-4.3); 23.3% of head (19.1%-32.4% of head).

Snout: 3.3 (2.3-4.2) ; 24.4% of head (17.9%-32.0% of head).

Distance from snout: to origin of dorsal fin 21.4 (17.6-25.8)
[27.9]; to origin of anal fin 84.8 (80.8-88.4) ; to origin of
ventral fin 42.1 (38.5-50.0).

Pre-anal length without head: 71 (66-76).

Dorsal fin: rays 6, rarely 5 or 7.

Anal fin: rays 10-13.

282 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Pectoral fin: rays 11-14.

Ventral fin: rays 7, rarely 6 or 8.

Vertebrae: 54-62.

Branchiostegal rays: 15-21.

Serial photophores: Ventral row: IP 8-11; PV 18-22; VAV
24-29; AC 9-13; Total L62] 63-70 [72]. Lateral row: OV
17-22; VAL 23-29; Total 42-49.

Body slender, elongate and compressed, its depth averaging
less than 1/12 of the standard length. Chin barbel present in
young, slender, tapered, flexible, becoming reduced to a short,
triangular stump in adults. Indo-Pacific populations have
slightly longer barbels than do Atlantic specimens, and appear
to retain the fully developed barbel longer.

Head 1/10 to 1/6 of standard length, bony ridges above eyes.
Eye round, its anterior-posterior diameter averaging almost
1/4 of head length, and a little shorter than snout. Snout
short, but generally slightly longer than eye. Subocular organ
present below anterior part of eye, deeply embedded. Postocu-
lar organ round, below posterior part of eye.

Mouth large, jaws almost equal to head. Premaxillaries with
four teeth, second longest, fourth tooth usually longer than third.
Second tooth, and sometimes third and fourth also, slightly
barbed. Maxillary longer than premaxillary, with numerous
small, oblique denticles on posterior half of its ventral margin,
entering gape. Mandible long and heavy, with five to nine
(usually six or seven) large teeth, of which the first is much
the longest, Behind these, near corner of mouth, 0 to 10 or more
tiny denticles (see Table IV).

Pectoral fins arising below posterior edge of opercular appar-
atus, low on body, fins of 11 to 14 rays. Origins of ventral fins
close together on midventral line, before middle of standard
length, fins normally of seven rays, rarely six or eight. Dorsal
with six (rarely five or seven) rays, its origin over the 5th to
8th OV photophore in Atlantic and Mediterranean populations,
as far posterior as the 10th OV photophore in some Indo-Pacific
individuals. Anal origin far behind dorsal, close to caudal, fin of
10 to 13 rays. Caudal fin deeply forked.

Sides of body with five longitudinal rows of scales, each scale
area marked off by the underlying pigment pattern in skin.
Each scale area with one to several small light organs, those of
the second row with one or two organs, those of the third row
with two, fourth row with one to three, most often two. In

MORROW : TAXONOMY OF CHAULIODUS 283

temperate Atlantic forms, the two organs of the second row are
usually unpigmented. In most of the Pacific, Indian and tropi-
cal Atlantic populations, the more ventral of the two organs
in the second row is usually pigmented. In some Pacific popula-
tions, the small unpigmented organ is reduced or absent, and
the same is true of many specimens from the Mediterranean Sea.
Large serial photophores located ventral to scale areas of fifth
row. SM organs vary in number from one population to an-
other, in most Atlantic forms, usually 1+2+1, in Indo-Pacific
forms basically 1+3+1 or 1+1+0, with many variations in the
number of small anterior and posterior organs.

Size. The largest specimen examined was 278 mm (11 inches)
in standard length. Zahl (1953:603) records one from the
Mediterranean of 350 mm, which appears to be the largest
known.

Relationships. Chauliodus sloani is the most basic form among
the modern species of the genus. At least two species, C. pam-
melas in the Indian Ocean, and C. schmidti in the Atlantic,
appear to have been derived directly from it. C. macouni of
the North Pacific and C. danae from the Atlantic, although un-
doubtedly derived from a sloani-like form, nevertheless show
rather more basic differences which suggest that their origins are
more remote than those of C. pammelas and G. schmidti. And
G. barbaius is obviously the farthest removed, morphologically
speaking, from the basic type.

Range. Of world-wide distribution in tropical and temperate
seas. In the Indo-Pacific, it is found as far south as approxi-
mately 42° S, and north to roughly 32°N. In the Atlantic, the
species has been recorded between 36°S and 57°N, with a single
record of a specimen washed up on the southeast coast of
Iceland at approximately 65 °N. It has also been recorded from
many stations in the Mediterranean Sea.

Discussion. The description and characterization of G. sloani
is not made easier by the distressing tendency of this form to
split into races which show greater or lesser morphological dif-
ferentiation, one from another. This has been thoroughly dem-
onstrated by Ege (1948). and is well shown in his illustrations
of the condition of the small body photophores in various
samples. Although the differences are not as constant as Ege
apparently believed, nevertheless, they do represent tendencies
within each population. There is, for example, a trend towards
pigmentation of both the small light organs in the second scale

284 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

row in the tropical forms, while in the populations from
temperate areas, these organs tend to be unpigmented. Yet at
the same time, greater or lesser numbers of individuals will be
found in any of these populations which do not agree with the
majority.

Perhaps even more striking is the change in the SM organs
with geography. The basic formula in the Indo-Pacific area
appears to be 1+3+0, that is, one unpigmented organ at the
anterior end of each section, folloAved by three pigmented organs.
But as one progresses eastwards through the Pacific, especially
in the Pacific Equatorial Water mass, this formula tends to-
wards 1+3+1 and finally 1+4+0. As far as this particular
feature is concerned, it suggests that C. barbatus, inhabiting the
eastern extreme of the Pacific Equatorial Water, is the end
result of whatever forces are at work here, for this species nor-
mally has five to seven small pigmented organs in each SM
series.

Movement away from the tropics, in a north-south direction,
is also reflected in changes in the SM organs. In northern areas
of the Pacific, between about 20°N and 32°N, the few specimens
available to us showed suggestions of reduction of the pigmenta-
tion of the last SM organ. In the temperate Atlantic, both north
and south, this condition is much more marked, with perhaps
the majority of specimens exhibiting a 1+2+1 condition.

A third characteristic is the length of the barbel, which is
shortest in the North Atlantic forms and longest in those from
the mid-Pacific. The Indian Ocean specimens are intermediate.
Also, the Atlantic C. slaani appear to lose their barbels at a
smaller size than do the Indian and Pacific ocean members of
the species.

All in all, these phenomena are open to at least two interpre-
tations. The first is that each population is more or less isolated
by the physico-chemical conditions of the water mass in which
it lives, and that the small differences observed from one popula-
tion to another are the result of reduced genetic interchange
between populations as compared to within populations. With-
out a doubt, this factor is operating but to what extent we can-
not tell.

It is also possible to interpret these slight changes in morphol-
ogy as representing the result of the direct response of the
organisms to physical and chemical characteristics of the partic-
ular water mass in which they live. We rather lean towards this

MORROW: TAXONOMY OF CHAULIODUS 285

interpretation, for we find that C. sloani, sensu stricto, occurs
in nearly all the water masses of the Indian, Pacific and Atlantic
oceans. Water mass boundaries do not appear, on the basis of
present information, to be serious barriers for this species.

As far as considering these populations as subspecies is con-
cerned, we have already shown that none of them meet the
modern criteria for such a division. There is greater difference,
for example, between the North Atlantic and Indian Ocean
populations of C. sloani than there is between the " secundus"
and "dannevigi" groups, yet no one has yet suggested — nor
do Ave do so now — that the two former groups should be desig-
nated as other than populations.
Synonyms and references :

Chauliodus sloani Bloeh and Schneider, Systema Ichthylogiae,
Berlin, 1801: 430 (type descr.) ; Cuvier, Regne Animal, Paris,
1836-1849: 232 (descr.); Cuvier and Valenciennes, Histoire
Naturelle des Poissons, Paris, Vol. 2k, 1849 : 382-389 (descr.,
synonymy) ; Canestrini, Fauna d 'Italia. Pesci., Milan, 1871-
1872: 221 (not seen); Doderlein, Atti Accad. Palermo, (2)
6, 1878-1879: 56 (name) ; Jordan and Gilbert, Bull. U.S. Nat.
Mus., 16, 1882: 285 (descr.) ; Facciola, Nat. sicil., 2, 1883: 188
(not seen) ; Jordan, Pep. U.S. Comm. Fish., (1885) 1887: 834
(name) ; Agassiz, Bull. Mus. Comp. Zool. Harvard, 15, 1888:
fig. 214 (illus.) ; Vaillant, Exped. Sci. Travailleur et Talisman,
1888: 102 (Morocco) ; Cams, Prodromus Faun. Med., Stutt-
gart, vol. 2, 1889-1893: 570 (name) ; Goode and Bean, Oceanic
ichthyology, Washington, 1895 : 96-97, fig. 115 (descr., illus.);
Garman, Mem. Mus. Comp. Zool. Harvard, 24, 1899 : 272-274,
PL K, fig. 3 (name, comp. with other spp.) ; Odon de Buen, Bol.
Soc. esp. Hist, nat., 2, 1902: 104-105 (descr.); Jordan and
Starks, Bull. U.S. Fish. Comm., 22 (1902) 1904: 579 (name) ;
Regan, Trans. Linn. Soc. Lond., (2) 22, Zool., 1908: 218 (N. of
Peros Banhos Atoll); AVeber, Siboga Exped. Fische, 1913:
12-13 (descr.) ; Weber and de Beaufort, Fishes Indo-Au-
stralian Archipelago, Leiden, vol. 2, 1913: 110, fig. 38 (descr.,
illus.) ; Sanzo, Mem. R. Accad. talassogr. ital., 39, 1914: 1-7,
figs. 1, 2 (descr. larvae) ; Gilbert, Proc. U.S. Nat. Mus., 48,
1915: 321 (name) ; Sanzo, B.C. Accad. Lincei, (5a) 27, 1918:
91-97 (descr. larvae) ; Gilchrist, Rep. Fish. Mar. Biol. Surv.
South Africa, No. 2, Spec. Rep. No. 3, 1921 (1922) : 42, 54
(occurrence S. Afr.) ; Jordan, Evermann and Clark, Rep. U.S.
Comm. Fish., (1928) 1930, App. X: 71 (name); Borodin,

286 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Bull. Mus. Comp. Zool. Harvard, 72 (3), 1931: 64 (name,
stomach contents) ; Fowler, Bull. Amer. Mus. Nat. Hist., 70,
1936: 219-221, 1199 (name, descr.) ; Bigelow and Schroeder,
Fish. Bull., U.S., 74 (vol. 53), 1953: 145-146 (descr.);
Tchernavin, Summary of the feeding mechanisms of a deep-
sea fish, Chauliodus sloani. Brit. Mus. (Nat. Hist.), London,
1953: 101 pp. (anat. head region in relation to feeding);
Munro, Marine and Freshwater Fishes of Ceylon, Canberra,
1955: 35 (Gulf of Manar, 500 fath.).

Chauliodus setinotus Bloch and Schneider, Systema Ichthyolo-
giae, Berlin, 1801 : 555, pi. 85 (type descr., illus., type locality
Mediterranean Sea) ; Bonaparte, Iconografia Fauna Italica,
III. Pesci., Roma, 1832-1841: fasc. XXX, No. 158, fig. 2 of
un-numbered plate (descr., illus.) ; Catologo Metodico, Napoli,
1846: 35 (name).

Esox stomias Shaw, General Zool., vol. 5 (1), London, 1804:
120, pi. Ill (descr.).

Stomias boa (Chauliodus schneideri) Cuvier, Regne Animal
Illus., Poissons, Paris, 1817: pi. 97, fig. 3 (illus., not seen).

Stomias schneideri Cuvier, Regne Animal, Paris, 1836-1849 : pi.
97, fig. 3 (illus., plate erroneously labelled Stomias boa).

Chauliodus schneideri Risso, Histoire Naturelle des Principales
Productions de l'Europe Meridionale, vol. Ill, Paris, 1826:
442, pi. 14, fig. 37 (not seen).

Leptodes sloanii Swainson, Nat. Hist. Classification Fishes, Am-
phibia, Reptiles, London, 1839: 298 (name).

Chauliodus sloanii (Giinther, Cat. Fish. Brit. Mus., vol. 5, London,
1864: 392 (descr.); Goode, Proc. U.S. Nat. Mus., 3, 1880:
483 (off So. New England) ; Vinciguerra, Ann. Mus. Stor.
nat. Genova, 22, 1885: 469-470 (descr.) ; Giinther, Handbuch
Ichthyol., Wien, 1886: 453, fig. 324 (not seen); Challenger
Rept., Zool. 22, 1887: 179 (New Guinea, Japan, Atlantic,
Bermuda, 565-2575 fath. water) ; Alcock, Ann. Mag. Nat,
Hist., (6) 4, 1889: 399 (Bay of Bengal, Gulf of Manar);
Collett, Bull. Soc. Zool. France, 15, 1890: 223 (Funchal) ;
Alcock, Ann. Mag. Nat, Hist., (6) 8, 1891: 127 (occurrence
Indian O., ova, pyloric caeca) ; Descriptive Cat. Indian Deep
Sea Fishes, Calcutta, 1899: 355 (not seen) ; LoBianco, Mitt,
Zool. Sta. Neapel, 15, 1902 : 419, 420, 422, 431 (juveniles, Gulf
of Naples); Regan, Ann. Mag. Nat. Hist. (9) 11, 1923: 614
(no. vert.).

Chauliodus sloanei Goode and Bean, Bull. Essex Inst., 11, 1879:

MORROW: TAXONOMY OF CHAULIODUS 287

22 (in stomach of cod, Georges Bank) ; Jordan and Ever-
mann, Bull. U.S. Nat, Mus., 47 (1), 1896: 585 (descr.) ; Rich-
ard, Bull. Mus. Oceanogr. Monaco, 41, 1905: 12 (name, N.
AtL, 0-3000 m) ; Brauer, Ergebn. Deutsche Tiefsee-Exped.
"Valdivia," 15, Syst. Teil, 1906:40-42, figs. 7-9 (descr.,
illus.) ; ibid., Anat. Teil, 1908: 48-59 (structure photophores),
176, (eyes) ; Zugmayer, Res. Cam}). Sci. Monaco, 35, 1911:
63-65 (descr.) ; Murray and Hjort, Depths of the Ocean, Mac-
millan, London, 1912: 603, fig. 454 (illus., name); 611
(name) ; Pappenheim, Deutsche Siidpolar Exped., 15, Zool. 7,
1914: 167 (N. Atlantic) ; Bierbaum, Z. wiss. Zool., Ill, 1914:
300-301 (structure ear) ; Roule, Res. Camp. Sci. Monaco, 52,
1919 : 26 (name, Med., Azores) ; Saemundsson, Vidensk. Medd.
naturh. Foren. kbh., 74, 1922: 177 (SE coast Iceland) ; Bar-
nard, Ann. S. Afr. Mus., 21, 1925: 141 (off E. London and
Durban); Regan and Trewavas, Dana Rept., No. 5, 1929:
32-34 (descr., range), fig. 24 (illus.) ; Beebe, Zoologica, N.Y.
12 (1), 1929: 10-12 (descr. young) ; Norman, Discovery Rept.,
2, 1930: 308 (S. AtL, partim) ; Roule and Angel, Res. Camp.
Sci. Monaco, 79, 1930: 24, pi. 1, fig. 23 (larvae, Canaries,
Azores, Med.) ; Res. Camp. Sci. Monaco, 86, 1933: 81 (name) ;
Beebe, Zoologica, N. Y., 22 (14), 1937: 201 (300-1000 fath.,
Bermuda) ; Parr, Bull. Bingham Oceanogr. Coll., 3 (7), 1937:
58 (Bahamas, Bermuda) ; Bertin, Bull. Mus. Hist. Nat.
Paris (2) 11, 1939: 382 (name); Norman, Sci. Rept. John
Murray Exped., 7 (1), 1939: 21 (name, Zanzibar, Maldives) ;
Nybelin, Goteborg. Vetensk. Samh. Handl., (B) 5 (16), 1948:
30 (NE Atl.) ; Ege, Dana Rept, No. 31, 1948 : 1-148 (classifica-
tion, phylogeny, zoogeogr.) ; Tchernavin, Proc. Zool. Soc. Lon-
don, US, 1949: 129-143 (cranial anat., mechanics swallow-
ing); Haffner, Systematic Zool., 1 (3), 1952: 112-133 (zoo-
geogr.); Grey, Fieldiana : Zool, 37, 1955: 277 (Bermuda,
stomach contents).

Chauliodus dentatus Garman, Mem. Mus. Comp. Zool. Harvard,
24, 1899: 273-274 (type descr, type locality Society Is, type
specimen Harvard Mus. Comp. Zool, No. 6597) ; Beebe,
Zoologica, N.Y, 12 (1), 1929: 12 (name).

Chauliodus dannevigi McCulloch, Biol. Res. "Endeavour," i
(4), 1916: 179-181, pi. LII (type descr.. illus, type locality
30 mi. S. of Cape Everard, Victoria, Australia, 180-240 fath.) ;
Regan and Trewavas, Dana Rept, No. 5, 1929: 32 (name).

288 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Chauliodus atlantis (partim) Barbour, Proc. New Engl. Zool.
Club, 19, 1942: 46, pi. XX (type descr., illus., type locality
21°46'N, 83°25'W, type specimen Harvard Mus. Comp. Zool.,
No. 35621).

Chauliodus danae Phillipps, Rec. Dominion Mus., 1 (1), 1942:
53-54 ( misidentification ) .

Chauliodus sloanei sloanei Ege, Dana Rept., No. 31, 1948 : 1-148
(classification, phylogeny, zoogeogr.) ; Haffner, Systematic
Zool., 1 (3), 1952: 112-133 (zoogeogr.); Koefoed, Rep. Sars
N. Atlantic Deep Sea Exped., 4 (2) (5), 1956: 18-19 (N. Atl.,
misc. discussion).

Chauliodus sloanei dannevigi Ege, Dana Rept., No. 31, 1948:
1-148 (classification, phylogeny, zoogeogr.) ; Haffner, Syste-
matic Zool., 1 (3), 1952: 112-133 (zoogeogr.).

Chauliodus sloanei secundus Ege, Dana Rept., No. 31, 1948:
1-148 (classification, phylogeny, zoogeogr.), 139 (type descr.,
type locality N. of Samoa Is., 300 m, type specimen Dana
Coll., No. St. 3588 (2)); Haffner, Systematic Zool., 1 (3),
1952: 112-133 (zoogeogr.).

REMARKS ON THE SYNONYMY

Chauliodus barbatus

The most easily distinguished species of the genus, it does not
appear to have been given any other name, nor have we found
any indication in the literature that it has been reported under
a mis-identification. Indeed, it is so different from the other
species of the genus that mis-identification hardly seems possible.

Chauliodus danae

Known only from the Atlantic Ocean, this is a well-defined
species, with a rather lower number of serial photophores than C.
sloani. Although Pappenheim (1914: 167) gave no character-
istics for his 22 mm specimen of "C. pammelas" from the North
Atlantic (24°41'N, 32°21'W), it seems probable that the speci-
men had rather fewer photophores than C. sloani, leading him
to name it as he did. This, together with the locality of capture,
strongly suggests that the specimen was actually C. danae.

As Ege (1948: 101) has shown, "C. danae" of Phillipps, from
Cook Strait, must belong to the "dannevigi" population of C.
sloani.

MORROW : TAXONOMY OF CHAULIODUS 289

We have examined the types and parat%ypes of Barbour's
(1942:46) C. atlantis, and find that the specimens "too small
to be determined with any certainty" belong to C. danae.

Chauliodus macouni

We have examined the type specimen of C. emmelas in the
U.S. National Museum, comparing it directly with the type and
other material of C. macouni in the same institution. There can
be no doubt that the two species are identical. The confusion
that has arisen over Jordan and Starks' (1904: 579) statement
that the head of C. emmelas is 7.5 in length, as compared with
"about one sixth" in C. macouni, is due entirely to the fact that
Jordan and Starks used total length, whereas Bean (1891: 44),
in describing C. macouni, used standard length. As we have
shown, there is no justification for retaining C. macouni as a
subspecies of C. sloani.

Chauliodus pammelas

Ege (1948: 101) points out that some specimens of "C.
sloani" described by Alcock (1899: 144) have only 61 photo-
phores in the ventral row, which is below the number known for
C. sloani. It is apparent, then, that these specimens must belong
to C. pammelas. Pappenheim's specimen of "C. pammelas"
from the Atlantic has already been discussed under C. danae. As
with C. macouni, we find sufficient divergence between C. pam-
melas and C. sloani to indicate that the former cannot be con-
sidered as a subspecies of the latter. We have therefore placed
C. s. pammelas as a full species.

Chauliodus schmidti

Ege (1948: 101) has pointed out that at least some of the
material reported as C. sloani by Norman (1930: 308) probably
belongs to C. schmidti, for "the two northernmost catching
places, . . . fall within an area off the west coast of Africa, where
Ch. sloanei sloanei is replaced by the new subspecies Ch. sloanei
schmidti."

We have shown above that this form must be regarded as a
species closely related to, but separate from C. sloani.

290 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ClIAULIODUS SLOANI

The various ancient names applied to this species are listed in
the synonymy, and would seem to present no problems worth
noting. A number of authors have followed Swainson (1839)
and Giinther (1864) in spelling sloani with two i's. There is no
justification for this, except that it was possibly the fashion in
the first half of the 19th century to use this form. Goode and
Bean (1879) used the spelling " sloanei," and Jordan and Ever-
mann (1896), evidently justifying their change to this emen-
dation, noted that the species was named for Sir Hans Sloane.
However, the spelling of patronymics in those days was a
casual matter, and Sir Hans himself was doubtless not entirely
certain of the correct spelling. We prefer to maintain Bloch
and Schneider's (1801) original orthography of "sloani."

We have examined the type specimen of Garman's C. denta-
tus (Harvard Mus. Comp. Zool., No. 6597), a species long con-
sidered distinct because of Garman's (1899) indication that
there were 14 AC photophores. Although the specimen is broken
in three pieces, nevertheless it is possible to fit the parts together
with a fair degree of precision. We counted 10 AC organs, with
probably one, possibly two missing from the break in the caudal
peduncle, for a maximum AC count of not more than 12. The
total count of the ventral row of photophores is either 69 or 70,
of the lateral row 48. These counts fall within the upper limits
observed for C. sloani, and since all other characteristics of the
specimen agree well with that species, we have placed C. denta-
tus Garman in the synonymy of C. sloani.

Examination of the type and paratype (Harvard Mus. Comp.
Zool., Nos. 35621 and 35615) of C. atlantis Barbour shows that
this species belongs in C. sloani. The smaller specimens accom-
panying the types have already been remarked on.

Chauliodus dannevigi McCulloch falls well within the normal
variation of C. sloani, according to McCulloch 's (1916: 179)
description, and is therefore included as a synonym. As Ege
(1948) showed, this form represents a more or less discrete
group, which Ege considered a subspecies. We find ourselves
unable to agree with this interpretation, considering dannevigi
to represent a population no more divergent than several others,
e.g. that from the Sulu Sea. The same is true of the form de-
scribed by Ege as C. sloanei secundus, as shown in the early
pages of the present paper.

MORROW: TAXONOMY OF CHAULIODUS 291

SUMMARY

Specimens of all known species of Chauliodus have been ex-
amined, including the types of C. barbatus, C. macouni, C.
emmclas, C. dentatus, and C. atlantis. As a result, only six
species of Chaidiodus, C. barbatus, C. danae, C. macouni, C.
pammelas, C. schmidti and C. sloani, are maintained. All other
species have been found to be synonyms. The several subspecies
proposed by Ege have been shown to be either fully recognizable
as species or else to be no more distinct than more or less discrete
populations of C. sloani.

LIST OF SPECIMENS1

Chaidiodus barbatus. Harvard Mus. Comp. Zool., Nos. 28489,
28490 (types); U.S. Nat. Mus., Nos. 148239, 151225; Zool.
Mus. Univ. Copenhagen, "Galathea" Coll., Nos. 12, 13, 14, 15;
Stanford Univ. Zool. Mus., No. 25314 ; Scripps Inst. Oceanogr.,
Nos. 52-363 (2), 52-390, 52-404, 55-244, 55-246, 55-258 (3),
55-265.

Chauliodus danae. Bingham Oceanogr. Coll., Nos. 2928 (6),
2929 (3), 2931, 2932, 2933, 2934 (2), 2935, 2936 (2), 2937,
2938 (5), 2939 (6), 2940 (4), 2941 (2), 3211 (3), 3766,
uncatalogued (8) ; Harvard Mus. Comp. Zool., Nos. 32277
(2), 34956 (9), 34983 (2), 35165, 35620 (3) ; U.S. Nat, Mus.,
Nos. 89914, 100336, 10056"), 117872, 117874 (2).

Chaidiodus macouni. U.S. Nat. Mus., Nos. 45372 (type), 51464
(type of C. emmelas), 53946, 77462, 125340, 132131 (2),
150088, 150089, 150090; U.S. Fish Wildl. Serv. Lab., Hono-
lulu, Nos. 1778 (3), 1780 (3), 1781, 1783; Stanford Univ. Zool.
Mus., Nos. 2000, 5259, 24950 (7) ; Scripps Inst, Oceanogr., Nos.
51-373 (4), 53-332 (3) ; Univ. Wash. Coll. Fish., uncatalogued
(2); Univ. Wash. Dept. Oceanogr., uncatalogued (57).

Chauliodus pammelas. British Museum (Natural History),
Nos. 1939.5.24.410, 1939.5.24.411, 1939.5.24.413, 1939.5.24.414,
1939.5.24.415, 1939.5.24.416, 1939.5.24.417.

Chauliodus schmidti. Marinbiol. Lab. Charlottenlund, Dana Coll.,
Nos. 3999 I, 4000 VI, 4000 X, 4003 V, 4003 VII (2), 4004 I
(3), 400.1 I, 4005 III, 4005 V, 4005 VII (7), 4006 I (8), 4006
II, 4007 T (3), 4007 VI (3), 4007 IX.

Chauliodus sloani. Bingham Oceanogr. Coll., Nos. 415 (2), 2915,
2916 (2), 2917 (2), 2918 (5), 2919, 2922 (4), 2924, 2925,

i Numbers in parentheses indicate number of specimens if more than one.

292 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

uncatalogued (4) ; Harvard Mus. Corap. Zool., Nos. 3903 (13),
26405 (2), 31607 (4), 31608 (6), 31609, 31610, 32274, 32275,
32276, 34949, 34952 (5), 34954 (2), 34978, 34982 (2), 35095,
35099, 36598, 39433 (5), 39726; Marinbiol. Lab., Charlotten-
hmd, Dana Coll., Nos. 3676 II (3), 3678 IV (4), 3678 VI,
3680 VI (7), 3680 VIII (4), 3681 I (4), 3682 I, 3683 I (3),
3683 IV (6), 3683 VI (5), 3683 VII, 3684 VI (2), 3685 I
(21), 3685 IX (3), 3686 VIII (4), 3735 I (6), 3735 II
(6), 3736 I (8), 3736 III (5), 3736 V (9), 3736 VI (2),
3737 I (18), 3737 II, 3738 I (8), 3738 II, 3738 III (2), 3739 I
(4), 3739 VI, 3739 VII, 3739 IX (2), 3740 I (4), 3740 II (2),
3925 II (3), 3933 III. 3949 I (2), 3953 I (2) ; as C. sloanei
dannevigi 3587 IX, 3601 I (2), 3613 VI, 3624 I, 3625 I, 3627
VI, 3630 II, 3631 II (2), 3631 III, 3639 I, 3640 I, 3640 III,
3651 I (3). 3651 VI, 3653 I, 3654 I, 3655 V (4), 3656 I,
3656 III, 3663 I, 3663 VIII (3), 3664 IV (3) ; as C. sloanei
secundus 3584 I, 3716 II (2), 3766 VIII (5), 3766 XIII, 3768
V, 3809 II (23), 3830 V (2), 3881 I (4), 3881 II (5), 3934
1 + VI+XI (3), 3934 II+VII+XII (5), 3934 III+VIII+XIII
(4), 3951 I, 3969 IV; Scripps Inst. Oceanogr., No. 52-404 (3) ;
U.S. Fish Wildl. Serv. Lab., Honolulu, 1782, 1917, 1918, 1919
(2), 1922, 1923, 1924 (2), 1926 (2), 2336, 2338, 2339 (2),
2340, 2341 ; U.S. Nat. Mus., Nos. 39218, 44402, 89915, 92246
(2), 117873, 131598, 143060, 158832, 159889, 163490 (2),
185590, 185591 (6), 185592 (2), 185593, 185594; Univ. Miami
Mar. Lab., Nos. 590., 716, 1623, 1632, 2004.

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Pappenheim, P.

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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE

Vol. 125, No. 10

A CONTRIBUTION TO THE BIOLOGY OF THE

GIGANTURIDAE, WITH DESCRIPTION OF A NEW

GENUS AND SPECIES

By Vladimir Walters

American Museum of Natural History

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

October, 1961

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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE

Vol. 125, No. 10

A CONTRIBUTION TO THE BIOLOGY OF THE

GIGANTURIDAE, WITH DESCRIPTION OF A NEW

GENUS AND SPECIES

By Vladimir Walters

American Museum of Natural History

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

October, 1961

No. 10 — A Contribution to the Biology of the Giganturidac,
with Description of a New Genus and Species1

By Vladimir Walters

INTRODUCTION

The morphology of the giganturid fishes is imperfectly known.
Brauer (1908) described the structure of the eye of Gigantura
indica, and Bierbaum (1914) worked out the structure of the
labyrinth of G. chuni. The osteology of G. vorax was studied
by Regan (1925). While preparing an account of the Giganturi-
dae for a forthcoming volume of "Fishes of the Western North
Atlantic" I found it necessary to investigate the anatomy of the
specimens, since many of their external features had been
destroyed.

William C. Schroeder and Mrs. Myvanwy Dick of the Museum
of Comparative Zoology, and James E. Morrow of the Bingham
Oceanographic Laboratory generously loaned their specimens of
Gigantura vorax. Through the kindness of Carl L. Hubbs and
Alfred Ebeling of the Scripps Institution of Oceanography,
two specimens of unidentified Pacific giganturids were bor-
rowed for comparison with G. vorax. I am grateful to N. B.
Marshall of the British Museum (Natural History) who ex-
amined the types of G. vorax Regan and G. gracilis Regan both
in the British Museum and in the Dana collections in Copen-
hagen. An expression of appreciation is due Giles W. Mead
of the Museum of Comparative Zoology who provided his manu-
script revisions of five families of iniomous fishes, and who for-
warded material belonging to various synodontoid genera. James
E. Bohlke of the Academy of Natural Sciences at Philadelphia,
Daniel M. Cohen of the U. S. Fish and Wildlife Service, Giles
W. Mead, and William A. Gosline of the University of Hawaii
critically read this manuscript. The sections dealing with the
nervous system, sense organs and sensory cues were read by
Lester R. Aronson of this Museum, and his suggestions and
criticisms are appreciated. Samuel B. McDowell, Jr. aided
in some of the dissections and interpretations of structure,

1 A more general account of the Giganturoidea will appear in Part 4, "Fishes
of the Western North Atlantic." Parts of these giganturid studies have been
supported by funds granted by the National Science Foundation (N.S.F. Grant
7123) to the Sears Foundation for Marine Research, Yale University.

298

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

»,

o
"3

'Si

Oil
• i-l

WALTERS: BIOLOGY OF THE GIGANTURIDAE

1299

including examinations of members of other groups which
formed the background for the phylogenetic assessment of the
Giganturidae. The drawings were made by Nina Williams and
Samuel B. McDowell, Jr.

Some of my observations (on other species) are at variance
with Regan's (1925) diagnosis of Gigantura and with his osteo-
logical account of G. vorax. For instance, none of my specimens
have vomerine teeth while G. vorax is said to have two vomerine
teeth, one behind the other; in G. vorax the scapula bears some

-pec
eth

eoc

Figure 2. Skull in dorsal aspect, A. Bathyleptus lime, holotype. B.
Gigantura vorax, American Museum of Natural History 20393. See legend
of Figure 3 for key to abbreviations.

of the fin-rays, the cleithrum extends the full length of the
girdle and there is no posterior coracoid process, while in my
material the scapula does not support any fin-rays, there is no
cleithrum, and the coracoid bears a posterior process. Marshall's
re-examination of Regan's material (both gracilis and vorax)
failed to disclose vomerine teeth, and the pectoral girdle, which
Regan illustrated, has either been discarded or lost. Marshall
also believes that the neurocranium of the type of G. vorax
more closely resembles the figures given below than it resembles
Regan's figures.

300

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

MORPHOLOGY

(Figures 2 to 7)

Visceral Arches. All that remains of the hyoid arch in Gigan-
tura and Bathyhptus are the hyomandibular and the quadrate;
the ventral elements of this arch and the branchiostegal rays are
absent. There do not seem to be any cartilages either supporting
or associated with the gills. In both genera the first gill slit
is the longest and lenticular, the second is shorter and elliptical,

soc

eth

den

Figure 3. Skull of Gigantura vorax in lateral aspect, composite. Ab-
breviations: an, angular; aut, autopalatine; hoc, basioceiptal; bsp, basi-
sphenoid; den, dentary; eoc, exoceipital; epg, eetopterygoid ; epi, epiotie ;
eth, mesethrnoid ; fr, frontal; liyo, hyomandibular; iop, interopercular;
lie, supraorbital lateralis canal; max, maxilla; mtp, metapterygoid ; n,
nasal ; op, opercular ; jxis, parasphenoid ; pec, preethmoid cornu ; phg,
preopercular-hyomandiljular gap ; pif, pineal foramen ; pop, preopercular ;
pro, prootic ; psp, pterosphenoid (alisphenoid) ; pto, pterotic; qu, quadrate;
soc, supraoccipital ; sop, subopercular; sphot, sphenotic; sub, suborbital
(?)•

the third is shorter and D-shaped, the fourth is very small and
circular, and there is no slit behind the hemibranch. In botb
genera the anterior set of pharyngeal teeth is in the roof of
the pharynx behind the last gill slit and anterior to the sus-
pensoria, while the posterior set is in the roof of the esophagus

WALTERS: BIOLOGY OF THE GIGANTURIDAE 301

medial to the kidneys. The posterior pharyngeal arch is the
most complete of the post-mandibular arches, consisting of a
backwardly directed pharyngobranchial cartilage (toothed),
epibranchial and ceratobranchial cartilages in line and sloping
downward and backward, and a forwardly-directed hypo-
branchial cartilage. There is no basibranchial cartilage. The
appearance of this arch is Z-shaped. The anterior set of pharyn-
geal teeth connects with the posterior set, and presumably repre-
sents the pharyngobranchial part of the last gill arch.1

Viscera. In both genera the kidneys are in the body wall be-
tween the epaxial and hypaxial musculature close behind the
pectoral girdle ; they are triangular and pale-colored with dis-
crete black spots. The anterior location of the kidneys indicates
that they may be pronephric, though they are not near the
heart. The excretory ducts follow retroperitoneal paths lateral
to the mesogasters and enter the urinary bladder beneath the
dorsal fin; the bladder discharges to the outside through a
urinogenital papilla. The ovaries are hollow sacs, fused pos-
teriorly, and in Gigantura they either discharge into the urinary
duct below the bladder or open to the outside so close to the
bladder that separate openings (if these exist) could not be
found in the material examined. The gross anatomy of the male
urinogenital system is unknown. In Bathyleptus the gonads are
missing from both specimens and the excretory duct cannot be
traced much beyond the rear of the stomach.

In both genera the cartilages of the pharyngeal arch embrace
the roof and sides of the esophagus ; a short muscle passes
from the lower end of the ceratobranchial and ends in a band
of muscle applied to the ventral face of the esophagus. The
cartilages have several bands of muscle between themselves and
the two sets of pharyngeal teeth. The posterior pharyngeals
have a set of retractor muscles which originate above the pyloric
valve in Bathyleptus, and above the posterior half of the stom-
ach in Gigantura; these retractors pass forward between the
two sheets of mesogaster and are inserted on the entire upper
surface of the posterior pharyngeals. In both genera the stomach
is a thick-walled blind pouch; when not distended with food

1 If we follow the system of E. Jarvik (1954 On the visceral skeleton of
Eusthenopteron with a discussion of the parasphenoid and the palatoquad-
rate of fishes. Kgl. Svensk. Veten. Hand. (4) 5 (1) : 1-104.) the anterior
pharyngeal set represents the infrapharyngobranchial and the posterior set
the suprapharyngobranchial elements of the same gill arch.

302

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

it is more than half the length of the body cavity and terminates
shortly in advance of the dorsal fin (in both specimens of
Bathyleptus the posterior end of the stomach is missing). The
inner lining, about one-quarter to one-third the total thickness
of the wall, is white and thrown into deep longitudinal folds.

Figure 4. Left pectoral girdle in lateral aspect. A. Bathyleptus lisac,
paratype. B. Gigantura vorax. Bingham Oceanographic Collection 3228.
Abbreviations: co, coraeoid; r\-r±, radials; pp, postcoracoid process; sr,
scapula.

This is surrounded by a thick layer of intensely black material.
External to the pigmented layer is the serosa. The stomach
hangs in the body cavity, suspended on each side by the meso-
gaster, which is muscular near the esophagus. The intestine,
which is colorless, leaves the pylorus near the midline, makes a
few small curves along which it receives ducts from the liver and

WALTERS: BIOLOGY OF THE GIGANTURIDAE

303

pancreas, ascends to the dorsal part of the body cavity, passes
caudad to the right of the mesogaster, turns ventrad between
the imfused lobes of the ovary, and terminates in an anal
papilla immediately in front of the urinogenital papilla. The
course of the intestine could not be followed beyond the torn
end of the stomach in Bathyleptus. There is one small pyloric
caecum. The astonishingly small orange-colored liver consists
of three lobes; in Bathyleptus the left-hand pair are joined
while in Gigantura the right-hand pair are joined. In Bathy-
leptus the liver lobes are horizontal; in Gigantura they are verti-
cal. In Bathyleptus the lobes are entire but in Gigantura the
right pair are subdivided into lobules.

ssii^^P^

~^m.

pig.

st L

Figure 5. Visceral relations. A. Bathyleptus llsae, holotype. B. Gigan-
tura vorax, composite drawing. Abbreviations: an, anus; hi, urinary blad-
der; es, esophagus; exc, excretory duct; int, intestine; fc, kidney; I, lobe
of liver; mpr, pharyngeal retractor muscle; o, ovary; ph, anterior and
posterior sets of pharyngeal teeth ; pig, deeply pigmented layer in stomach
wall; st. stomach.

Circulatory System. In Gigantura the pericardium is em-
braced by the liver, the ducts of Cuvier enter the short sinus
venosus beneath the middle of the ventricle, the truncus arterio-
sus is robust and conical, and three pairs of aortic arches arise
almost simultaneously from its anterior end. In Bathyleptus the

304

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

pericardium is anterior to the liver, the ducts of Cuvier enter
the long sinus venosus posterior to the ventricle, the truncus
arteriosus is small and bulbous, and three pairs of aortic arches
emerge from the short ventral aorta. In both genera the aortic
arches pass forward between the pharyngeal visceral arches be-
fore turning laterally and dorsally toward the gills. In both
genera the thyroid gland is anterior to the ventral aorta, and
apparently has no connection with the esophagus.

Figure 6. Heart and nearby structures in ventral aspect. A. Gigantura
vorax, Bingham Oceanographic Collection 3228. B. Bathyleptus lisae, holo-
type. Abbreviations : aa, aortic arches ; do, duct of Cuvier ; es, esophagus ;
I, lobe of liver; ph, pharyngeal arch; sv, sinus venosus; ta, truncus arterio-
sus; th, thyroid gland; v, ventricle; va, ventral aorta.

Brain, Cranial Nerves and Sense Organs. The brain of G.
vorax has been examined. In the paratype of B. lisae the head
and brain are crushed, but the outline of the brain is visible
through the roof of the holotype skull (cf. Brauer, 1908, table
42, fig. 18 for a figure of such a view of the brain of B. indicus).
From what can be seen, there do not appear to be any major
differences between the brains of the two genera. The forebrain
is very small. The enormous optic lobes are followed by the
enlarged cerebellum which does not override the optic lobes.

WALTERS : BIOLOGY OF THE GIGANTURIDAE

305

The walls of the medulla oblongata are thickened but there are
no enlarged sensory lobes. The stoutest cranial nerves are the
auditory (VIII), facial (VII), trigeminal (V), and optic (II) ;
the olfactory (I), oculomotor (III), trochlear (IV), and abdu-
cens ( VI ) are thread-like ; the glossopharyngeal ( IX ) and vagus
(X) are intermediate in stoutness. The pineal body is large
and applied to the inner surface of the frontals near the supra-
occipital; B. lisae has a foramen above the pineal, but in G.
vorax the frontals are separated by a cartilaginous wedge over
the pineal. The pineal stalk is remarkably long and robust ; at
the rear of the stalk is a pair of macroscopic habenular bodies
(microscopic in most teleosts). The hypophysis has a remark-
ably long and slender stalk.

The labyrinth of Gigantura chum was studied by Bierbaum
(1914) ; the sacculus is smaller than the utriculus. In a speci-
men in the American Museum of Natural History (No. 20393)
the plane of the horizontal semicircular canal parallels the longi-
tudinal axis of the head and body. The eye of B. indicus was
described in detail by Brauer (1908) ; my material of B. lisae
and G. vorax has not been studied in this regard. The olfactory
capsule is almost microscopic and the slender olfactory nerve
passes through the orbit dorsal and medial to the oblique muscles
(in both genera). The lateral line system is fairly well de-
veloped on the head although difficult to trace in entirety owing
to loss of skin; the infraorbital canal is particularly well de-
veloped in Gigantura, and is contained in flexible lightly-ossified

P,1

K1ZET

Figme 7. Brain and cranial nerves of Gigantura vorax, American Mu-
seum of Natural History 20393. Abbreviations: I, olfactory n. ; II, optic
n. ; III, oculomotor n. ; IV, trochlear n. ; V, trigeminal n. ; VI, abducens n. ;
VII, facial n. : VIII, auditory n. ; IX, glossopharyngeal n.; X, vagus n. ; ce,
cerebellum; fb, forebrain; ha, habenular body; hy, hypophysis; me, medulla
oblongata; op, optic lobe; pi, pineal body.

306 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

plates ( ? suborbitals) fringing the border of the mouth like a
transparent curtain. The vagus nerve was followed along the
trunk but a lateral line branch was not found ; there may not be
a trunk lateral line in these fishes.

BIOLOGY OF THE GIGANTURIDAE

General Considerations. The giganturids are mesopelagic
predators. All of the energy in the mesopelagic fauna is derived
from the photosynthetic processes of algae in the surface layers
of the sea; its downward flow is mediated principally through
the activities of diurnal vertical migrators which feed in the
upper levels and are in turn fed upon in the lower levels.
Lesser amounts of energy filter downward in the form of sinking
detritus. The deeper the level at which feeding takes place, or
in other words the greater the number of steps removed from the
primary surface production, the smaller is the amount of food
energy that is available for the biomass feeding at that particu-
lar level.1 The size and composition of the population at any
feeding level in the ocean is determined by the quantities of
food energy entering and of heat leaving that population per
time unit ; a state of dynamic equilibrium exists and if the flow
rate of energy out of the population can be reduced the result
would be an increase in the biomass. The flow of energy to the
physical surroundings (hence its escape from the biological sys-
tem) can be attenuated by reducing the rate of metabolism. Tt
is postulated that evolution in the mesopelagic and bathypelagic
biomasses has tended toward a reduction in the rate of loss of
heat energy to the physical surroundings through a reduction
in the metabolic rate.

The giganturids may have achieved metabolic economies in
several ways. By reducing the ossification of the skeleton, and
by retaining cartilage, the density of the fish is lowered. A
further reduction in density is achieved by flooding the sub-
dermal areas with a mesenchymal jelly. Having achieved a
density close to that of the surrounding water, the giganturid
has no need for a swimbladder and this structure is absent.

1 Although a mole of glucose contains about 675,000 calories, only 40 moles of
pyrophosphate with an energy content of about 400,000 calories are produced
when a mole of glucose is biologically oxidized to carbon dioxide and water.
In other words, some 275,000 calories per mole of glucose are lost from the
biomass at each link in the food chain.

WALTERS : BIOLOGY OF THE GIGANTURIDAE 307

Absence of the swimbladder eliminates the energy expenditure
necessary (in other fishes) for the maintenance of a gas bubble
at mesopelagial and bathypelagial pressures (see also Jones,
1957, p. 317).

On a wet weight basis, bathj'pelagic fishes which lack a swim-
bladder (Gonostoma elongatum, Xenodermichthys copei) have
2b' to 44 per cent of the protein content of shallow water species
with swimbladders (Ctenolabrus rupestris, Labrus bergylta)
(calculated from Denton and Marshall, 1958: table 1). This
suggests that the bathypelagic fishes have a metabolic level ap-
proximately one-third that of the coastal forms. The bathype-
lagic species were found to have dry weights 35 to 48 per cent
that of Ctenolabrus rupestris (recalculated by author) ; there-
fore, the bathypelagic forms have about two and one-half times
the water content of the coastal form, and, as Denton and
Marshall point out, the higher Avater content reduces the density
of the bathypelagic species. The ash of Ctenolabrus weighs about
1.8 to 2.4 times as much as the ash of Xenodermichthys, which
reflects the heavier and more extensive ossification of the coastal
species. Thus, Denton and Marshall's determinations are in
accord with my deductions concerning the Giganturidae.

Parr (1937) suggested that the animals of the deep sea consti-
tute a "rachitic fauna" since they are not exposed to sunlight
and live with little or no vitamin D ; this was offered to explain
the feeble ossification of deep-sea fishes. Marshall (1955: 324-
325) pointed out that abyssal benthic fishes are well ossified,
while some bathypelagic forms which are poorly ossified live
at levels not very far removed from the layers where vitamin D
is produced. The preceding paragraphs offer an alternative
explanation for the reduced ossification of some mesopelagic and
bathypelagic fishes ; it is likely that there are sufficient amounts
of vitamin D at all levels in the ocean, maintained through the
activities of vertical migrators.

It is my opinion that neoteny, indications of which are wide-
spread in many deep-sea groups, has been one of the major
avenues of evolution in the energy-poor deep-sea environment.
Neoteny eliminates part or all of the mainly catabolic phase of
life history, which is metabolically the most wasteful portion,
and by eliminating the morphogenesis of "adult" structures
neoteny may conserve additional amounts of energy for the
biomass.

308 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Sensory Cues. The enormous eyes, stout optic nerves, and
large optic lobes suggest that vision is important in the gigan-
turicl life history. In contrast olfaction is of minor importance ;
witness the minute olfactory capsule, thread-like olfactory
nerves and the minute forebrain. The other chemical sense, taste,
is evidently poorly developed since there are no strong sensory
enlargements in the medulla. The large cerebellum, stout audi-
tory nerve and small sacculus are indicative of a poor auditory
sense and a highly-developed sense of balance. The large tri-
geminal and facial nerves are partly associated with the cephalic
lateral-line system which is best developed along the border of
the upper jaw and apparently is of importance in feeding. The
enlarged pineal body, its robust stalk, and the large habenular
bodies may be associated with control of the response to varia-
tions in ambient light intensity. Iiasquin (1958) discussed in
detail the teleost pineal body and cited literature pertaining to
the role of the pineal in governing the response to light.

Denton and Warren (1957) calculated that teleosts should
be able to see daylight down to a depth of 1150 meters. This
is about the lower limit at which Gigantura chuni was collected
(660 fathoms) ; Baihyleptus lisae was taken in nets trawled at
1790 and 2140 fathoms but the actual depth of capture may
have been shallower. Weale (1955) noted that since in man
the threshold for light perception is 10 per cent lower for
binocular vision than for monocular vision, binocular vision
in deep-sea teleosts may double the visual response at illumina-
tion levels near the threshold for monocular vision. Thus gigan-
turids, which evidently possess binocular vision, should be able
to see their prey against an illuminated background throughout
their vertical range. They may hunt visually at light intensity
levels which are too low to enable the prey to see them well.
They may follow the deep scattering layers and prey upon the
luminous members of these layers, their silvery coloration allow-
ing them to blend inconspicuously with the illuminated back-
ground.

Swimming. The small number of vertebrae (about 30) indi-
cates that when a giganturid swims by producing lateral un-
dulatory movements of its body it employs the carangiform
rather than the anguilliform type of locomotion. The slight
side-to-side movement of the head during carangiform swimming
would aid in the search for food by broadening the horizontal
sweep of the visual field.

WALTERS : BIOLOGY OF THE GIGANTURIDAE 309

The plane of the horizontal semicircular canal being parallel
with the longitudinal axis of the body and the pineal body being
exposed through the roof of the skull indicate that giganturids
position themselves normal to the pull of gravity and normal to
the path of daylight, thus probably swimming horizontally.

The long lower caudal fin lobe may stabilize forward move-
ment in the yawing plane by lengthening the postanal part of
the fish so that the dorsal and anal fins lie in the first quarter of
the total length and pull rather than push the fish through the
water. The asymmetry of the caudal fin would pitch the snout
downward if the fin serves as a passive stabilizer and upward if
it is used to propel the fish. That the long lower caudal lobe
serves a passive, stabilizing role is indicated by the positions
of the pectoral fin bases which are somewhat higher in front
and would provide lift at the anterior end as the fish moves
forward, thereby offsetting the downpitch of the asymmetrical
caudal fin. The main propulsive force in swimming is appar-
ently derived from the muscular-based dorsal, anal and pectoral
fins.

Feeding. The gape of the mouth converges with the line of
sight. When prey is seen, the giganturid may slowly drift in
along its line of sight. When close enough so that a visual image
is in proper register on the retinas of both eyes, the fish may
pounce forward and seize its prey. Since the giganturid can-
not see what it swallows, the last -second location of the potential
food would be assumed by the infraorbital lateral line canals.

The sharp, recurved, depressible teeth indicate that once prey
has been grasped there is no escape ; and once it has been
seized, it must be swallowed. The prey is manipulated to the
rear of the mouth, perhaps through lurching and biting move-
ments of the giganturid. Eventually the anterior set of pharyn-
geal teeth is able to grasp it, and their stoking actions start the
food on its way down the esophagus. The posterior set of
pharyngeal teeth does most of the work in packing food into
the stomach, by rhythmic contractions of the powerful retractor
muscles. The pharyngeal retractors are antagonized by the
trunk musculature, which keeps the head and anterior trunk
rigid during swallowing (although in some preserved specimens
the pharyngeal retractors are contracted and the neck is dis-
torted). The pharyngeal visceral arch is used to maintain a firm
grip on the food while it is in the esophagus. In this way it
would quiet the struggles of the prey and thus prevent damage
to the giganturid 's heart and aortic arches, which are otherwise

310 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

unprotected. The mesogaster, which is muscular anteriorly, may
further aid in quieting the struggles of the prey.

Regan (1925) found a Chauliodus 140 mm. long in the stomach
of a Gigantura vorax 80 mm. long. A Gonostoma 86-mm. long
was found in the stomach of a 77-mm. Gigantura vorax (MCZ
35605). In both cases the tail and head of the food animal were
intact while its mid-body which, to judge by its position in
the stomach was swallowed first, was well-digested. These ob-
servations indicate that the giganturid requires a fairly long
time to swallow a large victim. Since a 65-mm. Gigantura vorax
(MCZ 40706) contained an intact 36-mm. Cyclothone sp. in its
stomach, we surmise that small prey can be swallowed relatively
quickly. In addition to respiratory problems which are dis-
cussed below, the giganturid is vulnerable to attack from other
predators while it is engaged in swallowing luminous prey. Per-
haps the giganturid packs as much as possible of the victim into
its stomach as quickly as possible and then closes its mouth over
the remainder which fits into the distensible, black-lined throat.
The thick deeply-pigmented layers of the distensible stomach
would conceal whatever light may be emitted by the victim
while it is being digested. When the food passes to the colorless
intestine it is presumably no longer luminescent.

The body cavity accommodates the enormous distention of the
stomach while a meal is being digested. The viscera are crowded
to the anterior and posterior ends of the body cavity, out of the
way of the stomach. The intestine runs dorsal to the stomach.
Without such an arrangement of its organs, the giganturid
would suffer intestinal strangulation and other discomforts with
each hearty meal.

Respiration. The small sizes of the second, third and fourth
gill slits and the absence of the gill flap from the upper end
of the gill cover indicate that giganturids do not respire in nor-
mal teleostean fashion. Regan (1925) suggested that the pec-
toral fins fan water into the gill chamber, but this is unlikely
since the gill flap would interfere while the portion which is not
covered by the gill flap is anterior to the pectoral fin. Hence the
giganturid probably respires by moving its suspensoria. When
the suspensoria expand, some water enters the gill chamber from
the pharynx via the first gill slit, when the fish is not feeding.
Water can also enter the gill chamber from the outside anterior
to the pectoral fin and gill flap. During inspiration the flexible
flap would be pressed against the body wall. Since the pectoral

WALTERS: BIOLOGY OF THE GIGANTURIDAE 311

fill base projects forward in the gill chamber over two sets of
gills, movements of the fin muscles could aid in mixing the water
in the chamber and bathing the gills. During expiration the
gill flap lifts away from the body wall and water leaves along
the entire length of the gill cover. The beating of the pectoral
fins wafts the water downward, backward, and away from the
gill chamber. While engaged in swallowing, the giganturid
would not obtain any oxygenated water via the pharynx ; all
would come from the outside.

SYSTEMATIC POSITION OF THE GIGANTURIDAE

Brauer (1901, 1906) compared Gigantura (and Bathyleptus)
with the original description and figure of Stylephorus chor-
datus, later shown to be an allotriognath. Regan's (1925)
derivation of the Giganturidae from the Synodontidae is dis-
cussed below. Berg (1940) named the order Giganturiformes,
following Regan in part. Tchernavin (1947a) corrected Berg's
diagnosis of the order and compared Gigantura with the Lyo-
meri without success because the anatomy of Gigantura was not
well enough known. Fowler (1936, 1958) combined Gigantura
and Stylephorus in one suborder; I have examined Stylephorus
chordatus and cannot agree witli Fowler. Bertin and Aram-
bourg (1958) placed the Giganturiformes between the Syn-
branchiformes (sic) and Saecopharyngiformes (= Lyomeri)
without presenting any new information.

Adult giganturids possess certain features which are associ-
ated with early developmental stages in other teleost groups : the
kidney is anteriorly placed ; the fin-rays are not segmented ;
copious amounts of mesenchymal jelly underlie the skin ; scales
are absent ; the cartilaginous endocranium persists ; a vagal
portion of the lateralis system seems to be absent ; bones and
cartilages are absent from the gill supports ; there are no gill
rakers ; the dermal elements of the pectoral arch are missing ; and
the pectoral girdle has a postcoracoid process ; the ventral ele-
ments of the hyoid arch, including branchiostegal rays, are
missing ; various dermal bones of the skull are missing such as
parietals, opisthotics, and premaxillae. Thus, the Giganturidae
show evidence of having become neotenic or larvalized.

Gosline (1959) thought it strange that neoteny is rare in
teleosts. Actually it is not. Extreme cases of nearly complete
larvalization, such as in Schindleria, are seldom found, but
there is a broad transition zone from this one extreme to the

312 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

other in which the adult differs considerably from the immature
stages (e.g., eels). The Giganturidae belong in this transition
zone. The manifestation of neoteny, whether slight or consid-
erable, implies that relationships of the group cannot be assessed
unless the comparisons are confined to similarities or dissimi-
larities in ontogenetieally equivalent structures.

If those features which appear to be neotenic were momentar-
ily left out of consideration, the Giganturidae may be looked on
as mesopelagic synodontoids. ("Synodontoid" = belonging or
allied to the series of families Synodontidae-Bathysauridae-Har-
padontidae.) In fact, Regan (1925) derived the giganturids
from the synodontids, and there is a superficial resemblance
due mainly to the backward extension of the jaws in both
groups ; the suspensoria have become backwardly-directed,
evenly-curved arches and their caudad swing has rotated the
bones of the gill cover so that the subopercular excludes the
opercular from the hind margin of the gill flap, and the pharyn-
geal teeth now lie between the suspensoria. In the Harpadonti-
dae {Ilarpadon nehcreus, II. mierochir), the pectoral girdle
approaches the giganturid condition in that the posttemporal
connects to the skull by muscle instead of a bone-to-bone contact,
but the harpadontid girdle is still well developed and has all
elements. Regan's assessment of the relationships of the Gigan-
turidae might have been different had he noted the weakly-
ossified lamina which lies buried in the cheek above and behind
the "premaxilla"; apparently, this lamina was lost during the
preparation of Regan's material.

The feature which distinguishes the iniomous fishes from less
advanced teieosts is that the maxillae are excluded from the
gape by the premaxillae, which alone border the upper jaw.
The dentigerous bone in the upper jaw of the Giganturidae was
identified by Regan (1925) as the premaxilla, but by Berg
(1940) as the maxilla. Actually this bone might conceivably
be any one of the following: 1) premaxilla, 2) maxilla, 3) fused
premaxilla and maxilla, 4) autopalatine. The "premaxillae" of
Gigantura and Bathyleptus have the following characteristics :
they do not meet anteriorly, they attach anteriorly to the vomer
and mesethmoid and posteriorly to the ectopterygoids, and they
do not border on the posterior quarter of the upper jaw. Since
the "premaxillae" do not border the upper jaw in its entirety,
and since there is a separate ossification behind each "pre-
maxilla," possible identifications (2) and (3) seem unlikely.

WALTERS: BIOLOGY OF THE GIGANTURIDAE 313

Consequently, the "premaxilla" appears to be the autopalatine,
which would account for its anterior and posterior points of
attachment to the skull, and which would also explain the ap-
parent absence of palatines, as reported by Regan. If this in-
terpretation is correct, the premaxillae have been lost by the
Giganturidae. The small ossification behind the autopalatine is
not likely to be premaxilla owing to its position; one would not
expect the distal, free end of the bone to be retained while the
proximal, articular end is lost. The small ossification behind
the autopalatine is thus identified as the maxilla. This line of
reasoning leads to the conclusion that the Giganturidae are sub-
iniomes.

Non-identity of the Giganturidae with the iniomes, and iden-
tity with the sub-iniomes, is suggested also by the presence in
Bathylcptus and in Gigantura of a gap between the hyoman-
dibular and the preopercular ; a muscle passes downward from
the cranium through this gap to insert on the lower jaw. In
none of the synodontoids has such an arrangement been noted,
but there is a similar gap and muscle in Esox lucius and in
Argentina (placed by Berg, 1940, in the Clupeiformes). It
would be interesting to learn how many teleost groups have
this muscle, and whether it may be of any phylogenetic sig-
nificance.

The Giganturidae are considered here as representing an order
Giganturoidea characterized as follows : upper jaw bordered by
autopalatines and maxillae ; premaxillae absent ; maxillae not
attached to skull ; anterior myodome absent, posterior myodome
present ; orbitosphenoid, opisthotics, parietals absent ; suspen-
sorium directed backward as an evenly-curved arch ; a lower jaw
muscle passing through the preoperculo-hyomandibular gap :
pectoral girdle free of skull and lacking posttemporal, supra-
cleithrum, cleithrum and mesocoracoid ; pelvic girdle absent;
ventral parts of hyoid and branchial arches, including branchio-
stegal rays and symplectic, absent ; pseudobraneh, three holo-
branchs, one hemibranch ; no gill rakers ; pectoral fin base
projects into gill chamber.

The relationship of the Giganturoidea to other teleosts is
vague. It seems likely, however, that they belong somewhere
between the esocoid members of the Tsospondyli and the synodon-
toid members of the Iniomi.

Considering now the relationships of the Giganturoidea to
higher teleost groups, Regan (1925) suggested they may be a

314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

specialized offshoot from a line that led to the gulper eels, order
Lyomeri (=- Saccopharyngiformes). The magnificent studies of
Tchernavin (1947a, 1947b) on S a ceo pharynx and Eury pharynx
indicate many trenchant differences between the Lyomeri and
Giganturoidea ; Lyomeri have no supraoccipital, a bone which
is well developed in Giganturoidea ; Lyomeri have a special ab-
ductor mandibular muscle while Giganturoidea have the usual
depressor mandibulae muscle ; Lyomeri have five or six holo-
branchs while Giganturoidea have three; Lyomeri have a single
kidney, whereas Giganturoidea have paired kidneys; in Lyo-
meri the pectoral muscles originate on the pericardium, but in
Giganturoidea the pectoral girdle is not associated with the
heart. Harry (1952) pointed out further that the luminous
organ of the order Cetunculi (Cetomimus) is comparable only
with the luminous organ of Lyomeri. It is also noteworthy that
in Cetunculi (Cetomimus) as in Giganturoidea the pectoral
girdle is free of the skull (see above for pectoral girdle of
Tlarpadon) and the stomach is deeply pigmented while the
intestine is not (Parr, 1929). The skull of Cetomimus as illus-
trated by Parr (1929) is topologically not too different from the
Giganturoidea. In conclusion, the Giganturoidea may be a
specialized offshoot of a line that led from a sub-iuiomous group
such as the esocoids toward the synodontoid iniomes, and this
line later may have given rise to the Cetunculi and perhaps
eventuallv to the Lyomeri.

Family GIGANTUPJDAE

hi the following account, unless otherwise noted, statements
concerning Gigantura and G. vorax are based upon specimens
of G. vorax having the following standard lengths : 65 mm.
(Museum of Comparative Zoology No. 40706), 77 mm. (Mu-
seum of Comparative Zoology No. 35605), 116 mm. (American
Museum of Natural History No. 20393), 156 mm. (Bingham
Oceanographic Collection No. 3228). Full data for these will be
given in "Fishes of the Western North Atlantic," volume 4.

Diagnosis. Mesopelagic teleosts. Eyes large, tubular, directed
forward. Gape of mouth extends far behind eye. Pectoral fin
bases above gill openings. Pelvic fins absent. Dorsal fin behind
middle of body. No adipose fin. Anal fin either partly below
or entirely behind dorsal fin. Caudal fin forked and middle
rays of lower lobe lengthened. Fin rays unsegmented, some

WALTERS: BIOLOGY OF THE GIGANTURIDAE 315

branched. Anus beneath dorsal fin. Sharp, recurved, depres-
sible teeth on autopalatines, dentaries, ectopterygoids and upper
pharyngeals. A layer of mesenchymal jelly beneath skin. Scales
absent. Luminous organs absent. Coloration silvery.

Upper jaw bordered by autopalatines and maxillae. Auto-
palatines joined to vomer and mesethmoid anteriorly and to
ectopterygoids posteriorly. Maxillae not attached to skull, but
buried in upper lip behind autopalatines. Premaxillae absent.
Anterior myodome absent. Posterior myodome present. Basi-
sphenoid, pterosphenoid (= alisphenoid) present. Cartilaginous
endocranium persists beneath surface bones. Supraoccipital
meets frontals. Orbitosphenoid, opisthotics, and parietals ab-
sent. Suspensorium directed backward in an evenly-curved
arch. Anterior upper pharyngeals between suspensoria. Re-
tractor muscles of posterior upper pharyngeals originate on
vertebrae dorsal to stomach. Kami of lower jaw loosely con-
nected at symphysis. Throat a distensible membrane. Preoper-
cular a splint applied to lower two-thirds of suspensorium. A
gap between preopercular and hyomandibular, through which
a muscle passes. Opercular excluded from posterior edge of
gill flap by subopercular. Pectoral girdle not attached to skull:
posttemporal, supracleithrum, cleithrum and mesocoracoid ab-
sent. Pelvic girdle absent. Ventral parts of hyoid and branchial
arches, including branchiostegal rays and symplectic absent.
One post-hyoidean arch complete, supporting both sets of
pharyngeal teeth. Pseudobranch, three holobranchs, and hemi-
branch present ; no slit behind hemibranch. No gill rakers.
Pectoral fin base projects into gill chamber. Kidney close behind
pectoral girdle. Swimbladder absent. Vertebrae 30-31.

Two genera, Gigantura Brauer 1901 and Bathyleptus, de-
scribed below.

Bathyleptus,1 gen. nov.

Genotype. Bathyleptus lisae, sp. nov.

Diagnosis. Epiotics almost as large as supraoccipital. Pineal
foramen present. Preethmoid cornua present. Supraorbital
lateral line canal in a bony ridge running full length of each
frontal. Pectoral girdle cartilaginous, with scapula, coracoicl
and four radials. Pharyngeal retractor muscles short and

i From the Greek Bathos = deep: Leptos = slender; in reference to the
depth range and body shape.

816 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

slender, originating on fifth or sixth vertebra. Visceral arch
supporting pharyngeal tooth-plates connects with strap of
longitudinally striated muscle on ventral surface of esophagus,
(iill openings and three holobranchs extend onto throat. Sinus
venosus anterior to liver. Trunk vertebrae several times longer
than wide. Trunk pencil-shaped. Caudal peduncle depth equal
to or less than its width. Least caudal peduncle depth 29-51 in
standard length.

Comparison with Oigantura. The diagnosis of Gigantura which
is given below follows the form used for Bathyleptus. Since
the two accounts are mutually exclusive it is unnecessary to pre-
sent a side-by-side comparison of the genera.

Gigantura Brauer, 1901. The epiotics are considerably smaller
than the supraoccipital. There is no pineal foramen. Preeth-
moid cornua are absent. The supraorbital lateral line canal is
in a bony ridge on each frontal between the orbits but the ridges
do not extend much posterior to the orbits. The pectoral girdle
contains an ossified scapula, ossified coracoid, cartilaginous post-
coracoid process, and four ossified radials; the fin-rays are in-
serted on the radials. The pharyngeal retractor muscles are long
and robust, and originate on the 12th through 16th vertebrae.
The visceral arch which supports the pharyngeal tooth-plates
connects with a strap of transversely striated muscle on the
ventral surface of the esophagus. The gill openings and gills are
situated entirely above the jaws. The sinus venosus lies be-
tween the lobes of the liver. The trunk vertebrae are about as
long as they are wide. The trunk is heavy and compressed.
The caudal peduncle is at least 1.5 times deeper than it is wide.
The least depth of the caudal peduncle is contained 11 to 16
times in the standard length.

Species. Three nominal species are placed in Bathyleptus. See
the remarks on Gigantura inclica Brauer and Gigantura gracilis
Regan in the diagnosis of the following species.

Bathyleptus lisae, sp. nov.

Figure 1

Holotype. Scripps Institution of Oceanography No. H51-375 ;
September 15, 1951; eastern Pacific north of Hawaii at 31°54'-
31°36'N., 152°21'-152°03'W.: 1790 fathoms with 10-foot Isaacs-
Kidd midwater trawl; R/V HORIZON, R. L. Wisner; 168 mm.
standard length.

WALTERS : BIOLOGY OF THE GIGANTURIDAE 317

Paratype. Scripps Institution of Oceanography No. H51-377 ;
September 23-24, 1951 ; eastern Pacific north of Hawaii at
31°01'-31°09'N., 127°39'-127°24'W.; 2140 fathoms with 10-foot
Isaacs-Kidd midwater trawl; R/V HORIZON, R. L. Wisner ;
head crushed, about 127 mm. standard length.

Description. D 17-18 ; A U-15 ; P 37-38 ; C 17-18 ; vertebrae
ca. 30- 31; 10 enlarged autopalatine teeth on each side; 2
ectopterygoid teeth on each side. Pin counts include all ele-
ments ; italicized values are for the holotype.

Proportional measurements as per cent of the standard length
(values for holotype italicized) : head length to rear angle of
jaws 13.7 ; head width across sphenotics 6.0; autopalatine
length 8.7 ; body width at dorsal origin 3.0, 4.7 ; body width at
middle of caudal peduncle 2.1, 3.5; body depth at dorsal origin
3.6, 6.3 ; least depth of caudal peduncle 2.0, 2.4 ; snout to dorsal
origin 53.6, 54.8; snout to anal origin 72.4, 74.5; snout to pec-
toral beginning 11.6 ; length of dorsal base 16.1, 19.7 ; length of
anal base 8.7, 8.9; length of pectoral base 6.0.

Diagnosis. The new species somewhat resembles Gigantura
chum inclica Brauer (1901) of the Indian Ocean and Gigantura
gracilis Regan (1925) of the tropical Atlantic, both of which are
herewith transferred to Batliijleptus since they have extremely
shallow caudal peduncles. Bathyleptus gracilis may be a syn-
onym of B. indie its. The new species differs from the others as
follows: in lisae the anal fin is completely behind the dorsal fin
whereas in gracilis and indicus it begins beneath the rear portion
of the dorsal fin ; in lisae the anal fin base is longer than the
pectoral fin base whereas it is shorter than the pectoral fin base
in gracilis (condition unknown for indicus) ; lisae has 37-38
pectoral fin-rays versus 39-41 in gracilis and 40-43 in indicus.

The new species is named after my wife, Lisa.

LITERATURE CITED

i>ERG, L. S.

1940. Classification of fishes both recent and fossil. Trav. Inst. Zool.

Acad. Sci. U. R. S. S., 5(2): 87-517.
Bertin, L. and C. Arambourg

1958. Super-ordre des Teleosteens (Teleostei). In P. -P. Grasse (ed.),

Traite de Zoologie (Paris), 13(3): 2204-2500.

BlERBAUM, G.

1914. Untersuchungen fiber den Bau der Gehororgane von Tiefsee-
fisehen. Zeitschr. Wiss. Zool., Ill: 281-380, 2 pis., 17 figs.

318 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Brauer, A.

1901. Uber einige von der Valdivia-Expedition gesammelte Tiefsee-

iische und ihre Augen. Sitzber. Ges. Beforderung ges. Natur-
wiss. Marburg, No. 8: 115-130.

1900. Die Tiefsee-Fisehe. I. Systematischer Teil. Jena, 432 pp.
Wiss. Ergeb. deutsch. Tiefsee-Expedition "Valdivia" 1898-
1899, Vol. 15. (Giganturidae, pp. 310-312, Gigantura chiini,
pi. 1, tigs. 1-3.)

1908. II. Anatomiseher Teil. Ibid., 266 pp. (Gigantura indica, pp.
213-215, pi. 42, figs. 18-21; pi. 43, figs. 1-9.)
Denton, E. J. and N. B. Marshall

1958. The buoyancy of bathypelagic fishes without a gas-filled swim-
bladder. Jour. mar. biol. Assoc. United Kingdom, 37: 753-767.
Denton, E. J. and F. J. Warren

1957. The photosensitive pigments in the retinae of deep-sea fish.
Ibid., 36: 051-062.

Fowler, II. W.

1930. The marine fishes of West Africa. Bull. Amer. Mus. Nat. Hist.,
70(1): 605 pp.

1958. Some new taxonomic names of fishlike vertebrates. Notulae
Naturae, No. 310, 16 pp.

GOSLINE, W. A.

1959. Four new species, a new genus, and a new suborder of Hawaiian
fishes. Pacific Sci., 13: 67-77.

Harry, R, R.

11)52. Deep-sea fishes of the Bermuda Oceanographic Expeditions.
Families Cetomimidae and Bondeletiidae. Zoologk-a, 37: r>5-72.
Jones, F. R. H.

1957. The swimbladder. Chapter IV. In M. E. Brown (ed.) The
physiology of fishes. Volume II. Behavior. Academic Press,
Inc., New York.

Marshall, N. B.

1955. Studies of alepisauroid fishes. Discovery Reports, 27: 303-336.
Parr, A. E.

1929. A contribution to the osteology and classification of the orders
Iniomi and Xenoberyces. Occ. Pap. Bingham Oceanogr. Coll.,
No. 2, 45 pp.
1937. Concluding report on fishes. Bull. Bingham Oceanogr. Coll.,
3(7): 1-79.
Rasquin, Priscilla

1958. Studies in the control of pigment cells and light reactions in
recent teleost fishes. Part 1. Morphology of the pineal region.
Part 2. Reactions of the pigmentary system to hormonal stimu-
lation. Bull. Amer. Mus. Nat, Hist., 115(1): 1-68.

WALTERS: BIOLOGY OF THE GIGANTURIDAE 319

Regan, C. T.

1925. The fishes of the genus Gigantura, A. Brauer; based on
specimens collected in the Atlantic, by the 'Dana ' Expedi-
tions, 1920 22. Ann. Mag. Nat. Hist., (9)15: 53-59, 1 pi.,
4 figs.

TCHBRNAVIN, V. V.

1947a. Six specimens of Lyomeri in the British Museum (with notes

on the skeleton of Lyomeri). Jour. Linn. Soc. London,

Zoology, 41: 287-350.
1947b. Further notes on the structure of the bony fishes of the order

Lyomeri (Eurypharynx) . Ibid., 41: 377-393.
Whale, R. A.

1955. Binocular vision and deep-sea fish. Nature (London), 175

(4466) : 996.

Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE
Vol. 125, No. 11

VARIATION IN PABAMYXINE, WITH A REDESCRIP-
TION OF P. AT AMI DEAN AND P. SPBINGEBI BIGELOW

AND SCHROEDER

By R. Strahan and Y. Honma

Dept. of Zoology, University of Hong Kong, and Dept. of Biology,

University of Niigata, Japan

CAMBEIDGE, MASS., U.S.A.
PEINTED FOE THE MUSEUM

October, 1961

Publications Issued by or in Connection
with THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 125.

Breviora (octavo) 1952 — No. 145 is current.

Memoirs (quarto) 1864-1938 — Publication was terminated with
Vol. 55.

Johnsonia (quarto) 1941 — A publication of the Department of
Mollusks. Vol. 4, no. 40 is current.

Occasional Papers of the Department of Mollusks (octavo)
1945 — Vol. 2, no. 26 is current.

Proceedings of the New England Zoological Club (octavo)
1899-1948 — Published in connection with the Museum. Publication
terminated with Vol. 24.

Of the Peters ' ' Check List of Birds of the World, ' ' volumes 1-3, 4
and 6 are out of print; volumes 5, 7 and 9 are sold by the Museum,
and future volumes will be published under Museum auspices.

Publications of the
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The remaining stock of the scientific periodicals of the Boston
Society of Natural History has been transferred to the Museum of
Comparative Zoology for distribution.

Proceedings-- Volumes available: 3, 5, 6, 8, 11, 14-17, 20-22, 24-27,
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(1-3), $10.00; Volume 6, $5.00.

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is available.

Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE

Vol. 125, No. 11

VARIATION IN PABAMYXINE, WITH A REDESCRIP-
TION OF P. AT AMI DEAN AND P. SPRINGEBI BIGELOW

AND SCHROEDER

By R. Strahan and Y. Honma

Dept. of Zoology, University of Hong Kong, and Dept. of Biology,

University of Niigata, Japan

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

October, 1961

No. 11 — Variation in Paramyxine, with a Redeseription of P.
atami Dean and P. springeri Bigciow and Schroeder

By 11. Straiian and Y. Hon ma

Paramyxine is characterized by having six pairs of external
branchial apertures, crowded together towards the posterior end
of the branchial region. Consequently, the most anterior efferent
branchial ducts are several times the length of the posterior ones
(Fig. 2). These characters are sufficient to distinguish Para-
myxine from Myxine and related genera (Nemamyxine, Neo-
myxine, and Notomyxine) which have only one pair of external
branchial apertures, and from Eptatretus (= Bdellostoma) in
which the efferent branchial ducts are of equal length. Two
species have been described: P. atami Dean 1904, based on a
single specimen from 494 m off Cape Manazaru in Sagami Bay,
Honshu, Japan, and P. springeri Bigelow and Schroeder 1952,
based on three specimens taken at 400-600 m in the Gulf of
Mexico.

Matsubara (1937) threw doubts on the sufficiency of the de-
scription of the type genus and species by demonstrating a
considerable degree of variation in a small collection of myxinids
taken at about 275 m from Kumano Nada, S.B. of Kii Peninsula,
Honshu, Japan. These specimens did not conform completely
either to the generic or specific descriptions of P. atami, but they
were far closer to this species than to any other known forms
and Matsubara considered that his specimens were "referable
to the above species, although discrepant in several characters"
(p. 13). Matsubara was concerned mainly in demonstrating the
variation in the arrangement of the branchial apparatus and the
teeth, and has left no record of the relative body proportions of
his specimens.

Okada et al. (1948a, b) handled almost 1,000 specimens of a
myxinid from the commercial hag-fishery of Sado Strait, taken
at a depth of 60-160 m, off Niigata, N.W. Honshu, Japan. These
authors were more concerned with data of commercial value (sex
ratio, weight-length ratio, etc.) than with taxonomic features,
and, following Matsubara (1937), they regarded their specimens
as referable to P. atami even though, for example, the length
of the type specimen lay outside the range of their large sample.

Bigelow and Schroeder (1952) have described some features of
a specimen from Suruga Bay, S.E. Honshu, Japan (U.S. Nat.

324 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Mus. No. 161442) which they regard (again, in spite of some
discrepancies) as referable to P. atami Dean. On the basis of
this specimen, Dean's (1904) description of the type, and some
data from Matsubara (1937), these authors came to the conclu-
sion that their Gulf of Mexico specimens were sufficiently dif-
ferent from P atami to justify the erection of a new species
P. springeri Bigelow and Schroeder 1952.

Strahan and Honma (1960), in a brief account of the hag-
fishery of Sado Strait, tabulated the mean body proportions of
50 specimens from a sample of some 500 myxinids collected off
Teradomari, N.W. Honshu, Japan. We had overlooked Bigelow
and Schroeder 's (1952) paper and, somewhat uncritically, had
followed Okada et al. (1948a) in regarding our material as
typical P. atami. We wish now to re-examine the situation in
the light of more detailed measurements of some 140 specimens,
and with respect to the work of Bigelow and Schroeder.

The Teradomari Sample

The method of collection of the most recent sample has been
described by Strahan and Honma (1960). All the specimens
were taken on the night of 21/22 October, 1959, at a depth of
about 100 in, about 8 miles offshore of Teradomari-machi (near
Niigata City), N.W. Honshu, Japan. The animals were main-
tained in aquaria for 1-3 days and measured under urethane
anaesthesia. After measurement, the pituitary glands were re-
moved from the majority of the animals (this being the prime
object of the expedition) and the bodies discarded. Some 50
specimens, preserved in formalin, were retained for later exami-
nation.

Hagfish are remarkably lacking in taxonomic features suscep-
tible to quantitative formulation. External features that can be
studied quantitatively are limited to the over-all length, the num-
ber of gill apertures and their position on the body, the position
of the cloacal aperture, the number of slime glands and their
distribution with respect to the gills and the cloaca. The caudal
fin terminates ventrally at the posterior border of the cloaca, but
the anterior limit of the dorsal caudal fin is not clearly defined.
Similarly, the anterior limit of the ventral (anal) fin is difficult
to define. Internal features of taxonomic value are the number
and arrangement of the lingual teeth, and the number of gills.

Length. Specimens varied in length from 130 to 583 mm.
Hitherto, the longest recorded specimen was the type, 550 mm

STRAHAN AND HONMA: VARIATION IN PARAMYXINE

325

long. Matsubara's (1937) specimen was 475 mm long and Okada
et al. (1948) did not record any individuals longer than 500
mm. The length frequency of the Teradomari collection is illus-
trated in Figure 1 and, for comparison, the length frequency
histograms of Okada et al. (1948a) are included.

There is a slight difference in the mode of the length in
June, August, and October (respectively, 325 mm, 316 mm, and
343 mm ) but this may be without significance, particularly since
the later measurements are separated by more than a decade
from the earlier ones. However, there is a very suggestive in-
crease, from June to October, in the relative frequency of very
large individuals (400 mm and more in length).

50r

2 30

^ 20

~ IO

«!-

Q 40

JUNE .

1944

: iuVI

LL
O

DC 25

00 20

Z> isL

io •

■ AUGUST.

1944

rmm

OCTOBER. 1959

O SO IOO 150 200 250 300 350 400 4SO SOO 550

-49 -99 -149 -199 -249 -299 -349 -399 -449 -499 -549 -599

LENGTH (mm)

Fig. 1. Length-frequency histogram, P. atami taken in the vicinity of
Niigata, Japan. Data for June and August, 1944, taken from Okada et
al. (1948a). Shaded portions of histogram indicate relative numbers of
individuals over 400 mm in length.

326

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Okada et al. (1948b) concluded that spawning occurs off
Teradomari mainly in April and May, but that it may extend
into August. In late October we found many females with
apparently mature eggs. In view of the extreme uncertainty
regarding the spawning season (if any) of Myxine glatinosa and
Eptatretus stouti despite extensive and prolonged investigations
(see Bigelow and Schroeder, 1948; Conel, 1931), we feel that it
is better to leave the question open for the time being.

Branchial system. Dean (1904) defines the genus Paramyxine
entirely on the branchial system : ' ' Hyperotretes with branchial
apertures ventrad of sacs. Ectal branchial ducts of distinctly
unequal length, the most anterior several times the length of the
most posterior. The duct of the most anterior gill opening at the
surface opposite the fourth (or fifth) gill sac. Openings of
branchial ducts drawn close together and compressed trans-
versely, that of the ductus oesophagoeus, however, longitudi-
nally, the latter aperture of large size, its length equalling that
of the sum of the interspaces of several gills. Transverse con-
strictor muscles of the branchial region developed as a distinct
element in the region of the hindmost branchial sacs." His
diagnosis of the species, P. atami, adds that there are six gills,
that the ectal duct of the most anterior gill is three or more
times the length of the most posterior, that the base of the tongue

A B

2 cm

Fig. 2. A. Ventral view of head of P. atami, 460 mm long. B. Dissec-
tion of left branchial region of P. atami, 530 mm long. Abbreviations:
ao., ventral aorta; bif., bifurcation of aorta (right branch not shown);
cm., branchial constrictor muscles; d., efferent branchial duct; r/.a.l., first
gill aperture; g.s., first gill pouch; n.t.1,2, first and second nasal tentacles;
o.t.1,2, first and second oral tentacles; ph., pharynx; pli.il., pharyngo-
cutaneous duct; t., base of 'tongue' muscle.

STRAHAN AND IIONMA : VARIATION IN PARAMYXINE

327

muscles lies between the third pair of branchial sacs, and that
the gill apertures are white against a purplish skin.

The general arrangement of the gill-pouches and ducts in a
specimen from Teradomari is shown in Figure 2. It can be seen
that this agrees with Dean's description in many respects. How-
ever, as Matsubara (1937) noted, there is considerable varia-
tion in the position of the posterior end of the lingual muscula-
ture. In the individual depicted, this lies between the second
pair of gill pouches, but in different individuals its position
varies from the first to the fourth gills.

The position of the gill apertures was not recorded by Dean
(1904) nor by Matsubara (1937), but we agree with Bigelow
and Schroeder (1952) in regarding this as taxonomically sig-
nificant, although variable. In Tables 1 and 2 the distance from
the anterior extremity of the body to the first and to the sixth
gill slit is set out as a function of the total length of the animal.
There is a definite, though slight, tendency for the branchial

Table 1. Position of the 1st gill aperture in Paramyxine atami from Teradomari.

Based on 13? specimens.

Total length
(mm)

Snout to 1st pill aperture (?i of total length)

Arithmetic
mean - S.D.

21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 35

100 - 199

2 7 3

29.0 1 0.6

200 - 299

1238 18 4001001

28.8 i 1.8

300 - 399

4 6 15 10 3

28.1 t 1.0

400 - 499

1 0 0 G 1 6 12 10 15 2

27.7 1 1.6

500 - 599

2 2

26.5 1 0.5

All

1 0 0 0 2 14 23 35 50 12 0 0 1 0 0 1

28.2 I 1.6

Table 2. Position of the 6th gill aperture in Paramyxine atami from Teradomari.

Based on 139 specimens.

Total length

Snout to 6th Rill aperture {% of

total length)

Arithmetic
mean _ S.D.

(mm)

24 25 26 27 28 29 30 31 32 33 34

35 36 37 33 39

100 - 199

16 3 2

32.5 i 0.7

200 - 299

1 1 3 3 12 15 1

10 0 0 1

32.3 1 1.7

300 - 399

6 11 10 9 1

31.8 i 0.4

400 - 499

100012 3 9 IS 82

31.3 1 1.7

500 - 599

2 0 11

31.2 t 1.4

All

1 0 0 0 2 3 19 24 4 5 36 6

2 0 0 0 1

31.7 t 1.5

328

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

region to occupy a relatively more anterior position in larger
individuals than in shorter ones.

Matsubara (1937) and Strahan and Honma (1960) have
drawn attention to the very considerable variation in the ar-
rangement of the external branchial apertures, examples of
which are shown in Figure 3. The 12 individuals depicted consti-
tute a random sample which, although it is too small to indicate
the relative frequency of particular variations, indicates their
type and degree. Matsubara (1937) found only one of his 14
preserved specimens from S. E. Honshu in which the gill aper-
tures were arranged in regular linear series. The remainder
showed roughly the same degree of variation as those depicted
here. Bigelow and Schroeder (1952) had at their disposal only
one specimen of P. atami and this, like Dean's type specimen,

2 ¥:■ \

'-,■,■
•'■».

; >\

!?;»■ •

V *

'.', - *

■?;/,■'■

■■'• ,

I ., :

xV;-:

■ %

- *-:~:.i<

'■ ".>

•:%V

* ■" ;?

->■;-..* '

??* ■ » •

T^'

,c* • . "•

--*%•■>•'■■"■'

".■?

■ **\. ■ ■. ,v,:y

■•:

ffr

§;'.

r:.

.* .-

;'■■- •

-.■-

■ " -■■

t-iL'-'i '

> :

'#1

i .-1

WK. .

II f 1 •■ 1

0
1

3 4 5

1,1,1

12 !

Fig. 3. Variation in arrangement of gill apertures in P. atami. Camera
lucida drawings of formalin-preserved specimens.

had the gill openings regularly and evenly aligned. Comment-
ing upon Matsubara 's observations, these authors suggest that
the irregularities may be due to muscular contraction. We are
convinced that this is not so, since we have observed quite
irregular arrangements in heavily anaesthetised animals. On the
other hand, Ave agree with Matsubara (1937) in his suggestion
that the outline of the apertures is liable to change during

STRAHAN AND IIONMA : VARIATION IN PARAMYXINE 329

preservation. In the living animal, the apertures may be more
or less circular, triangular, or elongately oval, but the outline
is smoothly convex. After preservation in formalin, the more
elongate apertures may develop crenellations and become cres-
centic. However, if such specimens are placed for a time in
water, the turgor developing from osmotic inflow restores a
condition similar to that in life. The specimens depicted in
Figure 3 were so treated.

Dean (1904) regarded the transverse orientation as a generic
character. In common with Matsubara (1937) and Bigelow and
Schroeder (1952), we find that the variation is too great to sus-
tain this opinion. Similarly, we must reject Dean's criterion of
a longitudinally elongated aperture to the pharyngo-cutaneous
duct ("ductus oesophagoeus"). The pharyngo-cutaneous duct
and sixth branchial aperture communicate with a shallow com-
mon depression, partially covered by a more or less semicircular
fold of skin, and directed postero-mesially. The arrangement
is not particularly variable and we have found no example of a
markedly longitudinal depression. We can, however, confirm
that the gill apertures have unpigmented borders.

In a number of individuals (Fig. 3, Nos. 7 and 10) there are
only 5 gill apertures on the right-hand side of the body, and we
have seen one individual in which there were only 4. The left-
hand side is less variable in this respect, although one individual
was found with 7 apertures on this side, due to separate open-
ing of the 6th efferent branchial duct and the pharyngo-cutane-
ous duct. The existence of 5 apertures does not necessarily imply
that there are only 5 gill pouches. Dissection of three individu-
als with 5 apertures on the left side revealed 5 gill pouches in
one individual and 6 in the other two, the latter having one of
the apertures shared by two adjacent ducts.

The cloaca. The cloaca is slit-like, as in Myxine, and sur-
rounded by slightly swollen lateral lips. The ventral caudal
fin arises immediately posterior to the cloaca, and we have
therefore used the posterior border of the cloaca as a measurable
landmark for this structure. The ventral (anal) fin terminates
just anterior to the cloaca.

As can be seen from Table 3, the cloaca occupies a relatively
more posterior position in longer individuals. Since the bran-
chial region is relatively more anterior and the cloaca relatively
more posterior in longer specimens, the length of the abdomen
is quite markedly increased, and this is consequently a poor
taxonomic character.

330

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Slime glands. The slime glands, where they occur, are seg-
mentally arranged, one per myotome. The linear series begins at
the fourth or fifth myomere and extends back to the myomere
immediately in front of the 1st branchial aperture. A second
series begins at the level of the first postbranchial myomere and
continues to the region of the cloaca where there is a gap, rep-
resenting from one to four myomeres, after which a third series
continues for about ten myomeres up to the region of the caudal
heart. Thus, the slime glands may be divided into a pre-
branchial, an abdominal, and a caudal series.

Table 3. Position of the posterior border of the cloaca in Paramyxine ataini.
Teradomari sample. Based on 136 specimens.

Total length

Snout to posterior border of cloaca

-
Arithmetic

(mm)

(fc, of total length)

mean 1 S.I).

81 82 S3 34 85 86 87 83 89 90 91 92

100 - 199

6 0 5

86.9 t 1.0

200 - 299

1000178 17 3

87.2 + 1.4

300 - 399

3 1 7 16 9 2

87.9 + 1.2

400 - 499

7 19 15 3 1 1

88.5 t 1.0

500 - 599

10 3

88.5 t 0.8

All

1 0 0 0 4 14 23 57 30 5 1 1

87.8 t 1.1

The mean values for the three series and for the total number
are set out in Table 4. There is no correlation between the num-
ber of slime glands and the total length. This is perhaps to be

Table 4. No. of slime glands in left side of body of Paramyxine atami
from Teradomari. Based on 123 specimens.

Total length

Mean No. of Slime glands

Prebranchial

Abdominal

Caudal

Total No.

(mm)

Series

100 - 199

18.8 i 0.9

45.0 i 1.4

11.3 i 0.8

75.7 t 2.0

200 - 299

18.6 i 1.3

44.9 i 2.2

11.0 t 1.0

74.5 t 2.6

300 - 399

18.9 1 1.5

44.6 t 1.9

10.9 I 0„9

74.7 t 2.5

400 - 499

18.9 t 1.0

45.1 t 1.4

10.8 i 0.9

74.9 t 2.2

500 - 599

19.3 t 0.9

47'.3 2 0.9

11.5 t 1.1

77.7 ± 1.3

All

18.8 + 1.2

45.0 t 1.9 I 10.9 t 1.0

74.9 t 2.6

STRAHAN AND HONMA : VARIATION IN PARAMYXINE 331

expected if the number of slime glands is determined by the
number of myomeres. The total number of slime glands is fairly
constant, 95 per cent of the specimens examined having a total
number of slime glands falling within two standard deviations
of the arithmetic mean (71-79). The division of the glands into
three linear series is more variable. Of the sample of 123 indi-
viduals, only 3 have the arrangement suggested by the means
of each series (prebranchials 19: abdominals 45: caudals 11),
but 78 (63%) fall within one standard deviation of these means
(prebranchials 18-20: abdominals, 43-47; caudals 10-12), and
121 (98%) fall within two standard deviations (prebranchials,
16-21: abdominals 41-49: caudals 9-13).

Caudal fin. In our original description (Strahan and Ilonma,
1960, p. 29) we stated that "the dorsal caudal fin has no clearly-
definable origin, but may be said to arise at approximately the
level of the anterior border of the cloaca." However, Bigelow
and Schroeder (1952) state that their specimen of P. atami has
a dorso-caudal fin extending as far forward as the 6th pair of
gill apertures, although it is only a low ridge. These authors
also draw attention to Matsubara's (1937) illustration, which
shows a dorsal fin of about the same height as the ventral fin,
extending forwards to what they estimate as "one-third to one-
half the distance from the level of the cloacal pocket toward the
level of the 6th pair of gill openings," i.e., about 60 per cent of
the length of the body from the snout. Unlike the ventral (anal)
fin, the dorsal caudal fin is supported by fin-rays, and its limits
may be specified in terms of the distribution of these structures.
We skinned a small number of specimens and found a series of
dorsal fin rays extending anteriorly to about 65 per cent of the
body length from the tip of the head (see Table 5).

The difference between the apparent and real origin of the
dorsal caudal fin, i.e., the position of the most anterior fin-ray
and the point at which the fin becomes obvious as a thin mem-
brane distinct from the rounded contour of the body, is due to
several factors. The fin-rays anterior to the level of the cloaca
are short and inclined strongly towards the horizontal. Be-
tween them and the skin of the fin is considerable fatty con-
nective tissue which masks the discontinuity between the fin
and the body proper. It is unlikely that this part of the fin
is very much more prominent in life, for the skin here is thick
and tough.

332

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

The caudal fin is continuous around the tail and extends
forward ventrally as far as the posterior border of the cloaca.
The fin-rays in the ventral part of the caudal fin are unbranched,
in contrast to those in the dorsal and terminal part.

Table 5. Real and apparent anterior limits of the dorsal fin, based on 8
skinned specimens of Paramyxine atami.

Total length
(cm)

Apparent origin
Distance from
snout

Most anterior fin-ray
Distance from snout

Number

rays

(cm)

% Total length

(cm)

% Total length

139

112

191

160

130

278

225

175

291

241

186

295

245

183

310

254

207

353

295

228

355

300

227

Mean t S.D.

83 1 1

65 1 1

57 t 7

Bigelow and Schroeder (1952) found a difference in the con-
figuration of the ventral border of the caudal fin in P. atami and
P. springeri. They state (p. 4) that it is "nearly straight in all
three of the Gulf of Mexico specimens while it is pictured as
moderately convex in the two Japanese specimens of atami which
have been illustrated (Dean 1904, pi. 1, fig. 3; Matsubara 1937,
pi. 1, fig. A), likewise in the National Museum specimen of
atami (compare Fig. 5 with Fig. 6)." We do not find this to
be a specific character. In Figure 4, we have depicted the caudal
fins of ten randomly-selected individuals. These show slight
convexity and concavity, but are no less straight than that of
P. springeri (Fig. 4, no. 11). The posterior part of the fin has
a white border, one to two mm deep.

The ventral fin. In anaesthetized animals, it is difficult to
determine the anterior limit of the ventral (anal) fin. The
shrinkage caused by preservation renders the fin more con-
spicuous, but it does not permit very accurate localisation of
the junction between the thin fin and the thick ridge running
along the mid-ventral surface of the abdomen. In a sample of
42 preserved specimens, we estimate the anterior border of the

STRAHAN AND H0N1IA : VARIATION IN PARAMYXINE

333

\ I mm

1 ...... . XV

-^™^m"^*'~r ! ' ~~r~^— -

.-„,-,..„: ■.-"-*--__ .-

i "V

A -

■ r I ■ t- i ' i <

f ■ xJfV

■

0 1 2 3 4 5 cm

| i__l i I i I i I i I

Fig. 4. Contour of the caudal fin in Paramyxine. Nos. 1-10, P. atami,
camera lueida drawings of formalin-preserved material. No. 11, P. springeri
(after Bigelow and Schroeder, 1952). Note variation in the arrangement
of the slime glands.

334

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ventral fin to be situated 47.3 + 2.7 per cent of the total length
of the body from the anterior tip. This value ranges from
39 per cent to 54 per cent; but we find no correlation between
this and the length of the body.

The lingual teeth. The procedure for dissecting the pituitary
of the animal involved splitting the dental skeleton longitud-
inally. The teeth of the left side only were counted and thus
we have no data on the degree to which asymmetry may occur.
The teeth were counted under a binocular microscope, care
being taken to lift away the skin fold covering the outermost
teeth of each row. The frequency distribution of the various
combinations of teeth on the outer and inner combs (see Table
6) is much the same as that published in our earlier paper,
but is based on a larger number of specimens. It may be noted
that the range does not come near to including 13 outer : 12-13
inner, which is the dental formula of the type specimen of P.
at ami.

There is no correlation between total length and the number
of teeth.

Table 6. Arrangement of lingua! teeth in Paramyxinc
atami from Teradomari. Based on 109 individuals.

Number of teeth

Number of

Percentage of

Outer

Inner

row

individuals

Differences between specimens from the east and west coasts

of Japan

Records of the occurrence of Paramyxinc have been collated
by Strahan and Ilonma (1960). On the Pacific coast of Honshu
it has been recorded from Aomori Province (Matsubara, 1955),

STRAHAN AND HONMA : VARIATION IN PARAMYXINE

335

Ibaragi Prefecture (Asano, 1956), Sagami Bay (Dean, 1904),
Suruga Bay (Bigelow and Schroeder, 1952), Kii Peninsula
(Matsubara, 1937), and Kochi Prefecture (Kamohara, 1952).
On the east coast of Honshu, there are records from the vicinity
of Niigata (Okada et al., 1948a, b; Strahan and Honma, 1960),
and from San-in District (Mori, 1956). Mori (1952) includes
P. atami in a check-list of Korean fishes.

There seems to have been no attempt to compare these speci-
mens in detail with the type, except in the case of Bigelow and
Schroeder who, like Dean, were limited to one specimen. Since
the Teradomari sample differs in several characteristics from the
type and from the U.S. National Museum specimen, both of
which are from the east coast of Honshu, we tried to get a
sample of Paramyxine from the Pacific coast of Japan, but
have been able to obtain only five specimens, from Kanbara, in
Suruga Bay. These are similar in body proportions to the U.S.
National Museum specimen (also from Suruga Bay) and they
have been grouped together with it for comparison with the
Teradomari sample, the type, and the specimens from the
Gulf of Mexico (Table 7).

Table 7. Taxonomic characters of Paramyxine populations from the east and
west coasts of Honshu, and from the Gulf of Mexico.

Character

Western
Japan

Eastern
Japan

Gulf of
Mexico

(3)

Teradomorij
(120 - 140)

Suruga Bay
(6)

Sagami Bay
(1)

Snout to 1st gill aperture {% T.L.)
Snout to 6th gill aperture (£ T.L.)
Snout to poet border cloaca (,f° T.L.)
Ho. prebrar-chial slirae glands
No. abdominal slime glands
No. caudal slime glands
Total no. slime glands
Mean no. teeth, outer row
Mean no. teeth, inner row

23.2 ± 1.6

31.7 t 1.5

37.8 + 1.1

18.8 i 1.2
45.0 i. 1.9

10.9 t 1.0
74.9 t 2.6
10.0 + 0.7
10.0 t 0.6

26.6 Z 1.0
29.9 i 1.2
87.1 1 0.3
16.6 t 1.1
45-5 1 1.1
9.0 + 1.1
71.0 t 2.1
11.5 i 0.5
11.5 + 0.5

25.5
27.9
85.5

17-20 ?2

58 ?"

io -r

13
12.5

23.1 t °«4
26.6 + 1.0

85.2 + 1.7

17.3 t 1.7
52.3 1 5.5
12.2 t 1.0
86.0 i 5.6
13.5

11.5

i, Figures in brackets under place names indicate 3ize of sample.
2. Dean's two illustrations are not in agreement with each other.

With respect to all the characters set out in Table 7 (except
the number of abdominal slime glands), the specimens from
Suruga Bay are significantly different from the Teradomari
sample (difference between the means is of the order of three
times the sum of the standard error of the means). In spite of

336 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

the small size of the Suruga Bay sample, this indicates a biologi-
cal difference between the two populations, although this is
probably not so great as to warrant sub-specific distinction,
since the joint non-overlap (Mayr, Linsley and Usinger, 1953)
of the most divergent character (no. of lingual teeth) does not
exceed 89 per cent.

Data on the type specimen are rather unreliable. The only
measurement given by Dean (1904, p. 14) is the total length
(about 550 mm). Bigelow and Schroeder (1952) attempted to
derive the dimensions from Dean's figures, but the figures them-
selves are suspect, particularly with respect to the slime glands,
which Dean regarded as too variable to be of taxonomic value.
The measurements given for the type specimen in Table 7 are
calculated from Dean's (1904) text-figure 2 and plate I, fig. 3.
Insofar as any reliance can be placed on them, they indicate
that the branchial region is even more anterior than in the
Suruga Bay specimens, while the cloaca is even more anterior
than in the Teradomari sample. The number of abdominal slime
glands appears to be considerably greater in his specimen than
in other Japanese Paramyxine. Nevertheless, there is insufficient
evidence to separate this single specimen from others that we
have examined. The Teradomari sample included a specimen
with body proportions fairly similar to the type (250 mm long;
snout to 1st gill aperture, 26% ; snout to 6th gill aperture,
28% ; snout to posterior border of cloaca, 89% ; pre-branchial
slime glands, 16 ; abdominal slime glands, 51 ; caudal slime
glands, 11), so it seems that the type may be regarded as
atypical, but within the range of variation which has been
demonstrated.

The Validity of P. springeri

Once the type description is questioned, the validity of the
separation of P. springeri from it must also be reviewed. How-
ever, reference to Table 7 shows that the specimens from the
Gulf of Mexico differ from the Teradomari and Suruga Bay
populations even more than they do from the type specimen.
There is a significant difference between the means of all the
characters tabulated for the Teradomari and American forms,
and the percentage of non-overlap is high for many of the
characters (snout to 1st gill aperture, snout to 6th gill aperture,
total number of slime glands, and number of lingual teeth),
exceeding 96 per cent. This, in itself, is probably only enough

STRAHAN AND IIONMA : VARIATION IN PARAMYXINE 337

to justify separation into subspecies. There is, however, a
further difference between P. atami and P. springeri which per-
mits a clear separation. This is the presence in P. springeri and
the absence in P. atami of slime glands in the branchial region.
As mentioned above, the prebranchial series of slime glands in
P. atami is clearly separated from the abdominal series by a
gap in the region of the branchial apertures. In P. springeri
there is a short series of glands in the branchial region, running
parallel to, and on the ventro-mesial side of the line of the gill
apertures. In the three specimens of P. springeri which have so
far been described, the number of glands in the "branchial"
series varies from 3 to 6.

Discussion

Dean (1904, p. 18) states, "The wide range in the variational
characters in species of myxinoids has long been appreciated, in
the matter, for example, of the number of gills and 'teeth' and
in the proportions of body regions and fins. My own observa-
tions lead me to the belief that in the case of myxinoids it is
peculiarly necessary to base specific determinations upon the
average characters of as great a number of individuals as prac-
ticable." Similar views have been expressed by Ayers (1894,
fide Worthington) and Worthington (1905), who worked on the
Californian Eptatretus, and by Bigelow and Schroeder^ (1948)
in their review of the reputed species of Myxine. However,
no authors have hitherto presented a simple analysis of varia-
tion within a myxinid population, expressed in terms of mean
values of quantitative characters and the standard deviations
of these means. Without such treatment, and quantitative con-
sideration of the degree of overlap of different populations,
there is very little that can be usefully said on the subject,

Our analysis makes it clear that there is a significant differ-
ence between P. atami from Teradomari and P. atami from
Suruga Bay. From this, we have deduced the existence of an
eastern and a western form of the species. Proof of this would
require more widespread sampling, to determine, for instance,
whether the populations intergrade with each other around the
north and south coasts of Honshu, or whether there is discon-
tinuous variation. There is a possibility of a cline extending
from the Sea of Japan to the Gulf of Mexico. It is dangerous to
assume this on three samples, two of which are rather small, but
examination of Table 7 suggests that such a cline may involve

338 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

a progressive shortening of the prebranchial region and an
increase in the number of lingual teeth from west to east over
the range of the genus.

The type specimen of P. atami is incompletely described, and
the diagnosis of the genus and species has been shown to be at
fault in several respects. AYe therefore propose a further de-
scription of the genus and species, and a simplified description
of P. springeri. The latter is based entirely upon the excellent
description of Bigelow and Schroeder (1952) and differs from
it mainly in selection of those characters which we regard as
taxonomically significant.

Paramyxine Dean 1904

Myxiniformes characteristically with 6 gills with separate
efferent branchial ducts of distinctly unecpial length, the most
anterior being more than twice the length of the most posterior,
opening separately to the exterior, except for the most posterior
on the left side, which opens into the same external aperture as
the pharyngo-cutaneous duct. External branchial apertures
lying close together in the posterior third of the branchial region,
the series on the two sides converging rearwards in a linear or
irregular arrangement.

Rostral flap above nasal aperture wider than long, nasal aper-
ture relatively larger in diameter than in Myxinc. Distal ends
of second pair of oral tentacles bluntly rounded.

Dorsal caudal fin apparently originates at level of cloaca (but
fin rays may extend farther forward), continuous around pos-
terior tip of body to posterior border of cloaca, ventral edge of
fin more or less straight. Ventral (anal) fin extends from
anterior border of cloaca to about 40 per cent of the length of
the body from the snout.

Body darker on back and sides than ventrally, edges of caudal
and anal fin and tips of tentacles paler than surrounding skin.
Branchial apertures with white borders. Distinct pale patch
over region of eye.

Up to about 60 cm in length.

Paramyxine atami Dean 1904

Six gills (rarely five on right side), external branchial aper-
tures in two irregular (rarely regular) rows on ventral surface.
Snout to first gill aperture 28.2 ±1.6 per cent of total length

STRAHAN AND HONMA : VARIATION IN PARAMYXINE 339

(western form) or 26.6 ± 1.0 per cent (eastern form). Snout
to posterior border of cloaca 87.8 ± 1.1 per cent of total length
(western form) or 87.1 ± 0.3 (eastern form). Slime glands
18.8 ± 1.2 prebranchial, 45.0 ± 1.9 abdominal, 10.9 ± 1.0 caudal
(western form). Slime glands 16.6 ± 1.1 prebranchial, 45.5
±1.1 abdominal, 9.0 ± 1.1 caudal (eastern form). Total num-
ber slime glands 74.9 ± 2.6 (western form) or 71.0 ± 2.1
(eastern form). No slime glands in region of branchial aper-
tures. Lingual teeth, 10 — 0.7 outer, 10 ± 0.6 inner (western
form) 11.5 ± 0.5 outer, 11.5 — 0.5 inner (eastern form).

Color, purplish -brown dorsally and laterally, grey ventrally.
Length, up to 58 cm. Coastal waters of Honshu (Japan), Korea.

Paramyxine springeri Bigelow and Schroeder 1952
(Simplified Description for Comparison with P. atami)

Six gills, external branchial apertures in two rather regular
rows on ventral surface. Snout to 1st gill aperture 23.1 ± 0.4
per cent of total length. Snout to posterior border of cloaca
85.2 ± 1.7 per cent. Slime glands 17.3 ± 1.7 prebranchial, 52.3
± 5.5 abdominal, 12.2 ± 1.0 caudal. Total number of slime
glands 86.0 ± 5.6. Slime glands on each side, ventro-mesially to
external branchial apertures. Lingual teeth 13-14 outer, 11-12
inner. Color, greyish-brown. Gulf of Mexico.

Supplementary Note

Teng (1958) has recently described a new species, P. yangi,
eight specimens of which have been obtained from fish markets
in Kaohsiung, Taiwan, and are assumed to have been caught
in waters to the southwest of Taiwan. P. yangi is characterised
by having only 5 pairs of gills and external branchial apertures,
arranged irregularly as in P. atami. The head is longer than in
P. atami or P. springeri. A summary of the major character-
istics is given below.

Mean and
Standard Dev. Eange

Total length (T.L.) 229±6 mm (198-250 mm)

Snout to 1st branchial aperture (% T. L.) 31. 7 ±1.3 (28.9-32.9)

Snout to 5th branchial aperture (% T.L.) 33.3±1.0 (30.7-34.9)

Snout to post, border of cloaca (% T.L.) 86.0±1.8 (81.2-87.6)

No. prebranchial slime glands 18.1±1.0 (17-20)

Xo. abdominal slime glands 37.1±1.5 (35-40)

Xo. caudal slime glands 8.6±1.1 (8-10)

Total no. slime glands 63.8±2.8 (60-69)

340 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Key to species of Paramyxine

la. Five pairs of gills; snout to 5th branchial aperture about 33% of total
length; about 65 slime glands on each side; number of abdominal
slime glands approximately twice the number of prebranehial slime
glands; slime glands absent between 1st and 5th branchial apertures;
9-10 lingual teeth in internal row, 9-10 in external row; up to 250 mm
in length yanyi Teng, 1958. Taiwan.

lb. Six pairs of gills.

2a. Snout to 6th branchial aperture about 30% of total length ; about
75 slime glands on each side ; number of abdominal slime glands
approximately 2.5 times the number of prebranehial slime glands;
slime glands absent between 1st and 6th branchial apertures; 9-10
lingual teeth in internal row, 9-10 in external row ; up to 583 mm in
length atami Dean, 1904. Japan.

21>. Snout to 6th branchial aperture about 27% of total length; about
85 slime glands on each side; number of abdominal slime glands
approximately 3 times the number of prebranehial slime glands ;
3-6 pairs of slime glands between 1st and 6th branchial apertures ;
11-12 lingual teeth in internal row, 13-14 in external row; up to

590 mm in length

springeri Bigelow and Schroeder, 1952. Gulf of Mexico.

ACKNOWLEDGEMENTS

The authors wish to express their gratitude to the following:
Dr. N. Kuroda and the Kanbara Fisheries Co-operative Associa-
tion, for collection of specimens from Suruga Bay; the officials
of Teradomari-machi, for the use of the town aquarium ; the Uni-
versity of Niigata, for provision of equipment and facilities ; the
University of Hong Kong for a grant towards travelling ex-
penses (R.S.) ; and, in particular, the Japan Gas Chemical
Company, without whose generous aid the work would have
been impossible.

Part of the work described was financed by a Research Grant
from the University of Hong Kong.

SUMMARY

Variation in the taxonomic characters of Paramyxine atami
Dean from the eastern and western coasts of Japan is described.
These populations differ significantly (in the position of the
branchial region, the number of prebranehial and caudal slime
glands, and the number of lingual teeth), but not sufficiently to

STRAHAN AND HONMA : VARIATION IN PARAMYXINE 341

rank as subspecies. The type specimen is not typical of either
population and appears to be an unusual specimen. Certain
points in the original definitions of the genus and species are
invalid and further descriptions are given. P. springeri Bige-
low and Schroeder, from the Gulf of Mexico, is found to be
taxonomically distinct from P. atami, particularly in possessing
slime glands in the branchial region. A simplified diagnosis of
of P. springeri is given. A note on the recently described species,
P. yangi is appended, with a key for separation of the three
species of Paramyxine

REFERENCES

Asano, N.

1956. Vernacular names of fishes in Ibaragi Prefecture. Japan J.
Ichthyol. 5: 19-51 (in Japanese).
Ayers, H.

1894. Bdellostoma dombeyi Lac. Biol. Lectures at Woods Hole in
Summer session of 1903. Boston (Ginn & Co.).

BlGELOW, H. B. AND W. C. SCHROEDER

1948. Cyclostomes. In: Fishes of the Western North Atlantic. Sears
Found. Mar. Res., New Haven, Mem. 1 : 29-58.

1952. A new species of the cyclostome genus Paramyxine from the
Gulf of Mexico. Breviora, Mus. Comp. Zool., No. 8: 1-10.
Conel, J. L.

1931. The segmentation of the egg of the myxinoid, Bdellostoma
stouti, based on the drawings of the late Bashford Dean. In:
E. W. Gudger (ed.) The Bashford Dean Memorial volume,
Archaic Fishes: 45-101. New York (Arner. Mus. Nat. Hist.).
Dean, B.

1904. Notes on Japanese myxinoids, a new genus Paramyxine and a
new species Homea olcinoseana, reference also to their eggs.
J. Coll. Sci. Imp. Univ. Tokyo 19: 1-23.
Kamohara, T.

1952. Revised descriptions of the offshore bottom fishes of Tosa
Province, Shikoku, Japan. Rep. Kochi Univ. Nat. Sci. 3: 1-122.

1958. A catalogue of the fishes of Kochi Prefecture (Prov. Tosa),
Japan. Rep. Usa Mar. Biol. Stat. 5: 1-76.
Matsubara, K.

1937. Studies on the deep-sea fishes of Japan. III. On some remark-
able variations found in Paramyxine atami Dean, with special
reference to its taxonomy. J. Imp. Fish. Inst., Tokyo 32 : 13-15.

1955. Fish morphology and hierarchy. Tokyo (Ishizaki-shoten). (In
Japanese.)

342 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Mayr, E., E. G. Linsley and E. L. Usingee

1953. Methods and principles of systematic zoology. New York
(McGraw Hill).
Mori, T.

1952. Check list of the fishes of Korea. Mem. Hyogo Univ. Agrie.

1: 1-228.
1956. Fishes of the San-in District, including Oki Islands and ad-
jacent waters (Southern Japan Sea). Mem. Hyogo Univ. Agric.
2: 1-62. (In Japanese, with English summary.)
Okada, Y., K. Kuronuma and M. Tanaka

1948a. Studies on Paramyxine atami Dean, found in the Japan Sea,
near Niigata and Sado Island. I. Misc. Eep. Ees. Inst. Nat.
Eesour. 11: 7-10. (In Japanese.)
1948b. II. Misc. Eep. Ees. Inst. Nat. Eesour. 12: 17-20. (In Japanese,
with English summary.)
Strahan, E. and Y. Hon ma

1960. Notes on Paramyxine atami Dean (Fam. Myxinidae) and its
fishery in Sado Strait, Sea of Japan. Hong Kong Univ. Fish.
J. No. 3: 19-26.
Teng, F. T.

1958. A new species of cyclostome from Taiwan. Chinese Aquatic
Products (Chinese Fisheries), No. 66: 3-6. (In Chinese.)
Worthington, J.

1905. Contribution to our knowledge of the myxinoids. Amer. Nat.
39: 625-663.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 125, No. 12

ABYSSAL MOLLUSKS FROM THE SOUTH ATLANTIC

OCEAN

By Arthur H. Clarke, Jr.
National Museum of Canada

With Four Plates

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

October, 1961

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 125.

Breviora (octavo) 1952 — No. 145 is current.

Memoirs (quarto) 1864-1938 — Publication was terminated with
Vol. 55.

Johnsonia (quarto) 1941 — A publication of the Department of
Mollusks. Vol. 4, no. 40 is current.

Occasional Papers of the Department of Mollusks (octavo)
1945 — Vol. 2, no. 26 is current.

Proceedings of the New England Zoological Club (octavo)
1899-1948 — Published in connection with the Museum. Publication
terminated with Vol. 24.

Of the Peters "Check List of Birds of the World," volumes 1-3, 4
and 6 are out of print; volumes 5, 7 and 9 are sold by the Museum,
and future volumes will be published under Museum auspices.

Publications of the
Boston Society of Natural History

The remaining stock of the scientific periodicals of the Boston
Society of Natural History has been transferred to the Museum of
Comparative Zoology for distribution.

Proceedings-- Volumes available: 3, 5, 6, 8, 11, 14-17, 20-22, 24-27,
30-34, 37. $4.00 per volume.

Occasional Papers: Volume 2, $5.00; Volume 3, $4.00; Volume 4
(1-3), $10.00; Volume 6, $5.00.

Memoirs : Requests for some specific memoirs can be filled but no list
is available.

Bulletin of the Museum of Comparative Zoology

AT HARVAED COLLEGE

Vol. 125, No. 12

ABYSSAL MOLLUSKS FROM THE SOUTH ATLANTIC

OCEAN

By Arthur H. Clarke, Jr.
National Museum of Canada

With Four Plates

CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM

October, 1961

No. 12 — Abyssal Mollusks from the South Atlantic Ocean1-2
By Arthur H. Clarke, Jr.

INTRODUCTION

During the twelfth cruise of the Lamout Geological Observ-
atory research vessel VEMA (see Ewing and Heezen, 1956)
made in 1957, extensive biological and geological explorations
were carried out in the Argentine Basin and off the west coast
of Africa from the Agulhas Basin to the Guinea Basin. Six-
teen successful bottom trawls were made in these regions,
fourteen of which were from abyssal depths. In 1958, the
fourteenth cruise of the VEMA again allowed opportunities
to trawl in the South Atlantic, and eleven bottom samples were
taken from the area between South Georgia and the Cape of
Good Hope. Eight of these were also from abyssal depths. The
numerous mollusks collected are nearly all quite different from
North Atlantic forms, and many new species are present.
Several of these are described on the following pages.

A summary of information regarding the trawling stations
is given in the tables below. At station 12, a bottom trawl with
an opening 3 meters wide was employed but at each of the
other stations a 1 meter trawl was used. Both trawls were
equipped with a fine mesh nylon net which retains all benthic
animals larger than small foraminifera. At each station the trawl
was in contact with the bottom for approximately 30 minutes.

R/V VEMA Abyssal Trawling Stations in the
South Atlantic Ocean

L.G.O.
Station No.

Latitude

Longitude

Corrected Depth
(fathoms)

38°58.5' S

41° 45' W

2805

30°14.9'S

13°03' E

1703

28°25.2'S

8°28.5'E

2770

25°33' S

12°27' E

1660

23°00' S

08°11' E

2262

22°58.5' S

07°00' E

1510

22°41' S

03°16' E

2767

5°53.5' S

09°51.5' E

1675

6°19.3'S

08°18.5'E

2193

i This paper is part of a doctoral thesis accepted by HarTard University in
May, 1960.

2 Contribution No. 503, Laniont Geological Observatory (Columbia University),
Palisades, New York.

346

BULLETIN: :

MUSEUM

OP COMPARATIVE

ZOOLOGY

4°23.9

00°18'

2315

55°19'

37°57'

2030

55°29'

37°57'

2054

56°37'

34°38'

1902

56°43'

27°41'

1497

57°39'

13°37'

2064

45°34'

06°02'

2507

41°03'

07°49'

2711

36°34'

14°08'

2670

34°35'

17°31'

993

29°44'

37°15'

2727

212

47°57.5

48°03'

3334

Additional information is available for some of the stations,
and since it may be significant it is included below.

Supplementary Information on R/V VEMA
Stations in the South Atlantic Ocean

L. G. O.

Station No.

Surface
Sediment

Bottom
Temp. (°C.)

Oxygen
(ml/L.)

Dominant Animal
Groups

red clay

crustaceans,
mollusks

foraminifera

2.43

4.8

crustaceans,
echinoderms, worms

white clay +
Mn nodules

worms

foraminifera

echinoderms,

crustaceans,

nematodes

foraminifera

1.35

foraminifera

red clay

2.5

3.4

greenish lutite

worms, crustaceans

red clay

0.76

worms, crustaceans

foraminifera

crustaceans

In addition to giving the localities in terms of distance and
direction from selected points on the land, on the following
pages ocean basins are also cited. Basin terminology follows
that used by Sverdrup, Johnson, and Fleming (1942). Such
a procedure is considered useful in the present study, and it is
hoped that it will be adopted generally in other works dealing
with the mollusks of the open sea.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 347

ACKNOWLEDGMENTS

Prof. Maurice Ewing generously provided the opportunity to
study the mollusks on which this report is based. Much addi-
tional material for comparison was freely made available by
Drs. William J. Clench and Harald A. Rehder. Dr. Clench and
Dr. Ruth D. Turner also read the manuscript. The research
was conducted at Harvard University and was supported by
the Lamont Geological Observatory by means of Rockefeller
Foundation gifts (RF. No. 57076 and No. 54087) to that insti-
tution. The National Museum of Canada provided clerical and
photographic assistance during preparation of the final manu-
script. All of this aid is sincerely appreciated.

SYSTEMATIC SECTION

Class GASTROPODA

Order ARCHAEOGASTROPODA

Family FISSURELLIDAE

Genus PlTNCTURELLA Lowe 1827

Type species: Patella noachina Linne, by monotypy.

Subgenus FlSSURISEPTA Seguenza 1863

Type species: Puncturella (Fissurisepta) papillosa Seguenza,
subsequent designation, Woodring, 1928.

Puncturella (Fissurisepta) agulhasae, new species

Plate 1, figure 3 ; Plate 2, figure 9

Shell small, about % inch in length, conical, base oval ; fissure
apical and elliptical ; sculpture reticular ; periostracum brown ;
and septum vertical. Base width about 88 per cent of the length,
regularly ovate except slightly flattened anteriorly and finely
crenulated by the radial ribs. Anterior and posterior slopes
straight. Fissure apical, 1.0 mm. long, 0.9 mm. wide, and
elliptical except flattened posteriorly where it intersects the
apex of the septum. Fissure placed about 33 per cent of the
distance from anterior to posterior. Sculpture reticulate, con-
sisting of numerous, flat-topped, radiating costae and concentric
cords. Costae about 50 near the apex and increasing to about
140 near the base. Intercostal spaces slightly narrower than

348 bulletin: museum of comparative zoology

the costae and frequently bisected by radial threads which widen
and become costae as they descend toward the base. Concentric
cords about 90, not crossing the costae. Periostracum light
brown, present chiefly in the intercostal spaces between tbe
cords. Interior of shell whitish, glossy, and exhibiting the ex-
ternal sculpturing. Septum thin, vertical, originating at the
posterior edge of the fissure, slightly convex, extending about
Ys the distance to the base, and dividing the interior into two
approximately equal halves.

length width height

holotype 8.0 mm. 7.5 mm. 5.5 mm.

Types. The holotype is in the Museum of Comparative Zool-
ogy, no. 22495.3, from R/V VEMA station 51, Agulhas Basin
about 1000 miles southwest of Capetown, South Africa, in 2507
fathoms. It was alive when collected and is the only specimen
obtained.

Remarks. This species clearly belongs in Fissurisepta but it
is apparently entirely distinct from all other species. In general
shape it resembles P. (F.) tenuicola Dall 1927 (figured in John-
sonia, 2, pi. 64, figs. 4-6) from 294 fathoms off Cumberland
Island, Georgia, but that species is nearly smooth externally, is
striated internally, and the septum is directed posteriorly. The
other known species in the subgenus are more dissimilar.

Fissurisepta is primarily an archibenthal group with wide but
scattered distribution in the subtropical portion of the North
Atlantic, and the present species represents a sizable extension
of the geographic and bathymetric range of the subgenus. For
an excellent review of the genus Puncturella in the Western
Atlantic see Johnsonia, 2: 116-148 (1947).

Records. Known only from the type locality.

Family SEGUENZIIDAE
Genus SEGUEXZIA "Jeffreys" Seguenza 1876

Type species: Seguenzia formosa Jeffreys, subsequent desig-
nation, Harris, 1897.

Verrill (1884) erected a separate family (Seguenziidae) for
Seguenzia and Basilissa (Watson, 1879) and because of its radu-
lar characteristics placed it next to the Strombidae (Mesogastro-
poda). Other authors (Dall, 1889a, 1927b; Woodring, 1928:
Johnson, 1934) retained the group as a family but did not agree
on its relationships. Thiele (1925, 1931) placed Seguenzia in

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 349

the Trochidae (Archaeogastropoda) under the subfamily Mar-
garitinae. This action was followed uncritically by Wenz
(1938).

The characteristic morphology of Seguenzia sets these archi-
benthal and abyssal gastropods apart from all others. They
resemble members of the archaeogastropod family Trochidae in
shell shape and in the possession of nacre, but the radula ap-
proaches the typical taenioglossate type of mesogastropods, and
the aperture is characterized by having one well developed pos-
terior sinus and one or two anterior ones. The present writer
therefore follows earlier authors in regarding Seguenzia as be-
longing to a separate family, the Seguenziidae. Its place in
the system is questionable, but for the present, it is left next
to the Trochidae following Johnson, 1934. Basilissa probably
belongs here also.

According to Wenz (1938 : 277) the oldest fossils of Seguenzia
are from the Eocene. There are also morphological similarities
between Seguenzia and the Ordovician to Devonian genus Pro-
turritella Koken 1889 figured by Wenz (1938, fig. 436 b, c, e)
and Knight (1941, pi. 39, fig. 5 a-c). Knight (personal com-
munication) concludes that the resemblance is probably super-
ficial.

In addition to the uncertainties regarding the systematic
status and position of Seguenzia, difficulties often arise in de-
fining the morphological limits of the included species. The
following quotation from Dall (1889a: 269) illustrates this
condition in S. formosa Jeffreys (= S. monocingulata [Seg.]).

"In examining the specimens of Seguenzia ... I find myself
in a dilemma. Either each separate individual is to be regarded
as a species, or the variability of the shells is very great. Per-
sistent study of the specimens has convinced me that the latter
is the true solution, and that the most evident characters, such
as the umbilicus (in some adult specimens) may be present or
absent; that the number of spiral threads, their strength and
sharpness on the basal disk, are entirely inconstant, and, while
in the typical formosa the ridge next to the suture is waved
or granulate, in many it is perfectly plain."

Similar variation is seen in other species of Seguenzia and
minor differences probably cannot be used to separate species.
It is with such a liberal attitude that the following specimens
have been identified, several of which are slightly different from
the type specimens or original figures with which they were
compared, but sufficiently close to be considered conspecific.

350 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Seguenzia Antarctica Thiele 1925
Plate 3, figure 10

Three specimens were found which seem to fit this species,
one each at station 18 (2262 fathoms, near the northern edge
of the Cape Basin and approximately 400 miles west of Walvis
Bay, South-West Africa), station 50 (2064 fathoms, near the
southern end of the Mid-Atlantic Ridge in the Atlantic Indian
Antarctic Basin), and station 53 (2670 fathoms, Cape Basin,
about 300 miles southwest of Capetown, South Africa). The
species is truly abyssal; the only other recorded specimen being
Thiele 's type collected from approximately 2520 fathoms at
63°16.5' S, 57°51' E in the eastern end of the Atlantic Indian
Antarctic Basin.

Seguenzia eritima Verrill 1884
Plate 3, figure 6

One specimen collected at station 12 (2805 fathoms, mid-
Argentine Basin, about 100 miles east-southeast of Buenos
Aires) and two at station 18 (2262 fathoms, north edge of Cape
Basin) seem to be nearer to eritima than to any other described
species. 8. eritima was recorded by Verrill from 1290 to 2033
fathoms south of Marthas Vineyard, Massachusetts. It is pos-
sible that the specimens here reported represent another, closely
related species but they are too immature and worn to describe.

Seguenzia elegans Jeffreys 1876
Plate 2, figure 7 ; Plate 3, figure 5

Two specimens, one from station 12 (2805 fathoms, Argentine
Basin) and one from station 18 (2262 fathoms, Cape Basin)
agree rather closely with specimens of elegans in the Jeffreys
Collection now in the United States National Museum (no.
181660). 8. elegans was described from between 740 and 1095
fathoms off Portugal. The specimens also somewhat resemble
8. orientalis Thiele 1925 from off East Africa in 379 fathoms,
but are apparently closer to elegans than to any other species.

Seguenzia carinata Jeffreys 1876
Plate 4, figure 6

Four specimens of this species were found, three at station
14 (1703 fathoms, Cape Basin, about 400 miles northwest of
Capetown, South Africa) and one at station 50 (2064 fathoms,

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 351

near the south end of the Mid-Atlantic Ridge, Atlantic Indian
Antarctic Basin). The identifications here are more certain
than in the case of any of the other seguenzias found. 8. cari-
nata has been recorded from depths ranging from 675 to 2199
fathoms at various localities in the North and South Atlantic,
but apparently not previously from the basins herein reported.

Seguenzia louiseae, new species
Plate 4, figure 4

Shell small (2.4 mm. high), trochiform, strongly carinate,
narrowly umbilicate, and fragile. Color white to slightly yel-
lowish. Whorls five, with a sharp, prominent, finely undulate,
peripheral carina and a second, large, slightly less angular
carina between the periphery and the suture. On the upper
whorls the latter carina appears to carry two parallel threads
on its blade. Peripheral carina visible only on the body whorl.
Basal disc with about twelve narrow additional carinae, the
most prominent being the outer carina and that surrounding
the umbilicus. Finer sculpturing on the spire consisting of one
fine revolving thread below the suture and two to four fine
revolving threads below the central carina. Fine, evenly spaced,
longitudinal, sigmoid threads are also present on the top of
the whorls and on the basal disc, gradually becoming finer on
the latter as they approach the umbilicus. These threads paral-
lel the lines of growth and indicate the presence of a well de-
veloped anterior and posterior sinus when the aperture is un-
damaged. Aperture iridescent within, subrhomboid, irregular,
and expanded where it intersects the carinae. Columella slightly
curved and extending in a tooth-like projection. Umbilicus deep
and narrow, about one-twelfth the width of the shell. Nuclear
whorl smooth, of medium size, paucispiral and slightly bulbous.
Operculum thin, light yellow, and ear shaped.

height (mm.)

width (mm.)

holotype,

station

(dead)

2.4

2.7

paratype,

< <

1 1

(dead)

2.3

2.2 1

< <

i i

1 1

(dead)

1.7

1 1

(alive)

1.6

< <

1 1

C <

(alive)

0.8

0.9

Types. The holotype and four paratypes are from R/Y
VEMA biology station 51 (2507 fathoms, Agulhas Basin, about

i Last whorl broken away.

352 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1-450 miles southwest of Capetown, South Africa). Two addi-
tional paratypes are from station 18 (2262 fathoms, northern
end of Cape Basin, about 400 miles west of Walvis Bay, South
West Africa). The holotype is no. 224951 and the paratypes
from station 51 are no. 224952 in the Museum of Comparative
Zoology. Paratypes from station 18 are no. 4739 in the Na-
tional Museum of Canada.

Remarks. In general shape, thickness, and in possession of an
acute, nearly blade-like peripheral carina, S. louiseae more
closely resembles 8. carinata Jeffreys than any other species.
The carina between the periphery and the suture is much
heavier than in Jeffreys' figures of carinata (P.Z.S. 1885, pi.
5, figs. 3, 3a) however, and the sinuous axial threads, so prom-
inent in louiseae, are absent on carinata. Although louiseae is
apparently a very distinct species, it seems to be intermediate in
general structure between carinata and the more heavily sculp-
tured species of Seguenzia, e.g. ionica Watson and cost ulif era
Schepman.

I take pleasure in naming this species for my wife, Louise R.
Clarke.

Records. Known only from stations 51 and 18, cited above
under 'Types'.

Family CYCLOSTREMATIDAE
ABYSSOGYRA, new genus

Type species: Abyssogyra vemae, Clarke.

Shell small, planorbiform, weakly sculptured, and with a
multispiral, corneous operculum. Sculpturing limited to lines
of growth and two faintly developed, revolving carinae. Addi-
tional characters are those of the type species.

Abyssogyra lacks the heavy sculpturing and the beaded oper-
culum of Cyclostrcma, Marryat. It differs from Circulus Jef-
freys in not being longitudinally ribbed. Pseudomalaxis Fischer,
Omalaxis Deshayes, Zalipais Suter, Omalogyra Jeffreys, and
the several genera erected by Bush (1897) and by Pilsbry and
McGinty (1945-46) all differ in prominent characters from
Abyssogyra.

Abbott (1950) has fixed the identity of the problematical
Cyclostrema cavcellata of Marryat, the type species of Cyclo-
strema, and this has had the effect of restricting the genus to
include only the small, planorboid, heavily sculptured species
with beaded opercula occurring in shallow water in the tropics.

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 353

This has left many of the deep sea species formerly placed in
Cyclostrema without a proper genus. After a careful search of
the literature, it has become clear that such is the case with
Cyclostrema normani Dautzenberg and Fischer 1897. Since the
species described below is apparently congeneric with normani,
a new genus (i.e. Abyssogyra) is necessary to receive it.

Abyssogyra vemae, new species
Plate 3, figure 4

Shell minute (1.8 mm. wide), planorbiform, weakly sculp-
tured, and white in color. Periostracum brown, thin, and pres-
ent only in small patches. Whorls two, nearly circular in cross-
section. Suture deep. Spire depressed, projecting only slightly
above the body whorl. Aperture circular except in the parietal
region where it is nearly straight and slightly thickened. Outer
and inner lips thin and sharp. Umbilicus wide and extending
to the nuclear whorl. Sculpture consisting of lines of growth
and two low, medially located, revolving carinae, one located
on the dorsal side of the whorls and one located ventrally. Two
additional revolving carinae intersect the ends of the straight
parietal lip : the upper one borders the suture, the lower ascends
into the umbilicus. Nuclear portion bulbous, unsculptured, and
consisting of one-half whorl. Operculum thin, corneous, yel-
lowish gray, multispiral, and spirally ridged.

major diameter minor diameter height

holotype 1.8 mm. 1.3 mm. 0.9 mm.

Types. The holotype was collected at station 49 (1497 fath-
oms, 56°43' S, 27°41' W, Atlantic Indian Antarctic Basin, south
of Traverse Island, South Sandwich Islands). Only one speci-
men, a living one, was obtained. The holotype is in the Museum
of Comparative Zoology, no. 224962.

Remarks. As stated above, in shell characters this species
resembles a Cyclostrema without prominent sculpturing. It is
probably closest to Abyssogyra normani (Dautzenberg and
Fischer) but that species differs in having the aperture width
approximately one-third the width of the shell and the spiral
carinae very faintly developed, while in vemae, the aperture is
approximately one-half the width of the shell and the carinae
are rather well developed. It is also somewhat similar to
Homalogyra denticostata Jeffreys 1884, except that in that
species the spire is depressed below the level of the body whorl

354 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

and the concentric sculpturing is slightly sigmoid and more
prominent.

Records. Known only from the type locality.

Genus BROOKULA Iredale 1912

Type species: Brookula stibarochila Iredale 1912, original
designation.

Through the kindness of Dr. Donald F. McMichael of the
Australian Museum, Sydney, and Dr. Harald A. Rehder of the
United States National Museum, I have been able to examine the
holotypes of Brookula stibarochila Iredale 1912 (PL 1, fig. 5)
and Vetulonia galapagana Dall 1913 (PL 1, fig. 1), the type
species of the genera Brookula and Vetulonia, respectively.

The anatomy and radular characteristics of these two species
are unknown, but examination of the shells has led to the con-
clusion that their relationships are sufficiently distant to make it
desirable to place them in different subgenera. B. stibarochila,
a shallow water tropical species, is minute, umbilicate, covered
with axial ribs which are approximately vertical, has a beaded
aperture which is nearly parallel with the columella, and has
a white, glass-like shell. Vetulonia galapagana, an abyssal
species, is much larger, non -umbilicate (the umbilicus is neatly
filled with a callus), covered with oblique axial ribs, has a plain,
oblique (prosocline) aperture and a white, somewhat chalky
shell.

In addition, many of the archibenthal and abyssal species
formerly placed in Brookula appear to belong to a third group
distinct from Brookula (sensu strict o) and from Vetulonia. This
group is described below as a new subgenus and is given the
name Benthobrookula.

BENTHOBROOKULA, new subgenus

Type species: Brookula {Benthobrookula) exquisita Clarke,
original designation.

Shells small, umbilicate, sculptured with axial ribs which
are approximately vertical and with spiral threads or costae,
with a plain, nearly vertical aperture, a relatively large and
slightly bulbous protoconch, and with a white, opaque, solid
shell.

Benthobrookula differs from Brookula (sensu stricto) in the
following characters. Benthobrookula has a somewhat bulbous

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 355

and relatively large protoconch ; the aperture is plain, i.e. not
beaded or otherwise sculptured although it may be thickened;
the costae are narrow and symmetrical in cross-section; and
the shell is not glassy and not translucent. Brookula (sensu
stricto) has a small protoconch ; the aperture is beaded ; the costae
are flattened, rounded in front and blade-like behind; and
the shell is glassy and translucent and appears like a tropical
species which, of course, it is. In operculum characteristics, the
two subgenera are very similar.

From Vetulonia, Benthobrookula differs as follows. Vetulonia
has the umbilicus neatly filled with a callus, the ribs are
oblique, and the aperture is strongly prosocline. Benthobrookula
is openly umbilicate and the ribs and the aperture are ap-
proximately vertical. Anatomical studies are necessary in all
three of these groups, and the results of such studies may sup-
port or confound the division here proposed.

Thiele (1925: 57, 71) pointed out that much confusion existed
in the proper placement of the small shells previously assigned
to Cyclostrcma (e.g. Brookula, YUrinella, etc.) and proposed
three groups to include them : (1) Skeneidae in which the radula
has 4 or 5 lateral teeth, (2) Cyclostrematidae which has only one
lateral tooth and a few weak marginal teeth, and (3) Vitrinelli-
dae which is taenioglossate. The radula of Brookula powelli, n.
sp. (PI. 4, fig. 9) most clearly fits the Cyclostrematidae, not-
withstanding possible differences in interpretation concerning
which are lateral and which are marginal teeth.

Brookula (Benthobrookula) powelli, new species
Plate 3, figure 7 ; Plate 4, figures 1 and 9

Shell minute (2.3 mm. wide), trochiform, depressed, umbili-
cate, prominently sculptured, and grayish white. Whorls 2%,
convex, separated by a depressed suture and forming a slightly
obtuse spire. Sculpture consisting of numerous, rather heavy,
narrow, longitudinal ribs (40 on the body whorl and 32 on
the penultimate whorl of the holotype) and numerous, very
fine revolving threads (about 30 on the body whorl of the
holotype) which do not cross the ribs. The threads become
somewhat stronger on the base of the shell, but except for the
four cords which surround the umbilicus, the spiral striations
never approach the ribs in strength. The four cords around
the umbilicus are of approximately the same strength as the
ribs. Aperture complete, ovate, somewhat angled posteriorly

356 bulletin: museum of comparative zoology

and flattened where it is appressed to the preceding whorl.
Outer and inner lips slightly thickened. Umbilicus rather wide
and extending far up into the shell. Periostracum apparently
absent. Protoconch bulbous, smooth, and composed of one-
half whorl. Operculum thin, round, corneous, multispiral, and
yellowish. Radula (from a station 51 specimen) shown in
Plate 4, figure 9.

height (mm.)

width (mm.)

whorl

holotype, station 12

1.9

2.3

paratype, station 51

1.6

2.0

paratype, station 51

1.5

1.7

paratype, station 51

1.3

1.4

Types. The holotype containing the animal is from R/V
VEMA biology station 12 (2805 fathoms, mid-Argentine Basin,
about 1000 miles east -southeast of Buenos Aires, Argentina).
One paratype, living when collected, is from station 48 (1902
fathoms, about 100 miles southeast of South Georgia) and
thirteen paratypes, most of which contain the animal, are from
station 51 (2507 fathoms, about 1000 miles southwest of the
Cape of Good Hope). The holotype is no. 224960 in the
Museum of Comparative Zoology. Paratypes are in the Museum
of Comparative Zoology and the National Museum of Canada.

Remarks. Brookula (Benthobrookula) powelli is similar to
Brookula strebeli and B. pfefferi (both Powell, 1951, from off
South Georgia in 85 to 97 fathoms) in general appearance and
in the possession of umbilical cords, but those species are higher
than wide and exhibit 314 and 4% whorls, respectively, although
they are both smaller than the holotype of powelli which has
only 2% whorls. In addition, the axial sculpturing and the
spiral threads are finer on powelli.

The species is named for Dr. A. \V. B. Powell of the Auck-
land Museum, who has been for many years the leading author-
ity on Antarctic mollusks.

Records. Known only from the localities cited above under
'Types'.

Brookula (Benthobrookula) exquisita. new species
Plate 3, figure 8 ; Plate 4, figure 2

Shell minute (1.8 mm. wide), trochiform, depressed, umbili-
eate, heavily sculptured, and white. Whorls 2%, convex, sep-
arated by a depressed suture and forming a spire which is

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 357

produced at an angle of about 120°. Sculpture consisting of
numerous rather heavy, narrow, longitudinal ribs (28 on the
body whorl and 23 on the penultimate whorl of the holotype)
and many prominent revolving cords (17 on the body whorl
of the holotype), which give the shell a cancellated appearance.
The cords on the top of the whorls are weaker than the ribs, but
on the base, except for the three major cords surrounding the
umbilicus, the cords and ribs are of approximately the same
strength. The three cords near the umbilicus are more widely
spaced and are much heavier than the ribs. Aperture circular
except slightly flattened where it is appressed to the preceding
whorl and lightly crenulated by the three major basal cords.
Outer lip somewhat thickened, inner lip a little thinner. Um-
bilicus deep, wide at the base and narrowing as it ascends.
Periostracum apparently absent. Nuclear portion bulbous,
smooth, and composed of one-half whorl. Operculum thin,
round, corneous, multispiral, and yellowish.

height (mm.) width (mm.) whorls

holotype, station 47 1.5 1.8 2%

paratype, station 47 1.2 1.4 2%

Types. The holotype and one paratype, both living speci-
mens, were dredged at R/V VEMA Biology Station 47 (2054
fathoms, approximately 60 miles south of South Georgia). The
holotype is no. 225954 in the Museum of Comparative Zoology
and the paratype is no. 4742 in the National Museum of Canada.

Remarks. This species more closely resembles B. powelli than
any other species, but the spiral cords are very much heavier
in exquisita and the spire is markedly more obtuse. B. pfefferi
Powell (1951) and B. strebeli Powell (Joe. cit.) are somewhat
similar also, but in those species the spire is acute and the
spiral threads are very weak, nothing like the robust thickness
of the threads in this species.

Records. Known only from the type locality.

Brookula (Benthobrookula) lamonti, new species

Plate 4, figure 3

Shell minute (1.5 mm. wide)., trochiform, not depressed, sculp-
tured, umbilieate, and white. Whorls 314. convex, separated by
a sharply defined, depressed suture and forming a sHghtly
acute spire. Sculpture consisting of numerous, sharp, somewhat
elevated longitudinal ribs (25 on the body whorl and 22 on

358 bulletin: museum of comparative zoology

the penultimate whorl of the holotype) and many fine, low,
revolving threads (about 20 in the holotype) which do not cross
the ribs. The threads become a little stronger on the base of
the shell, but except for the two threads which border the um-
bilicus, they are much weaker than the ribs. The two cords near
the umbilicus are about the same strength as the ribs. Aperture
ovate-ear shaped, angled posteriorly and flattened where it is
appressed to the preceding whorl. Outer and inner lips thick-
ened. Umbilicus rather narrow and deep. Periostracum ap-
parently absent. Nuclear portion partly broken in the holotype
but apparently bulbous, smooth, and composed on one-half
whorl. Operculum thin, round, corneous, multispiral, and yel-
lowish.

height (mm.) width (mm.) whorls

holotoype, station 47 1.5 1.4 3~Vi

Types. The holotype, an unique specimen, was collected alive
at R/V VEMA biology station 47 (2054 fathoms, Scotia Sea
about 60 miles south of South Georgia). It is at the Museum
of Comparative Zoology, no. 225953.

Remarks. This species is grossly similar to B. pfefferi Powell
(1951) and B. strebeli Powell (loc. cit.) (which are so similar
to each other that they appear to represent the same species),
but lamonti has a much thicker lip than either of those species,
the aperture is angular posteriorly while in pfefferi and strebeli
it is not, and the umbilical area in lamonti is more open and
quite differently sculptured. From poivelli and exquisita, de-
scribed above, lamonti may be differentiated by its slightly acute
spire, its heavier lip, its less prominent ribs, and its more chalky
appearance.

This species is named for the Lamont Geological Observatory,
the sponsor of the expeditions which led to the discovery of
the mollusks here described.

Records. Only one specimen is known. See under 'Types'.

Brookula (Benthobrookula) capensis, new species

Plate 1, figure 4

Shell small (3.4 mm. wide), trochiform, not depressed, heavily
sculptured, white on the ribs and light brown between them.
Spire turreted and acute. Whorls approximately 3 to 4 (nuclear
portion broken away), convex, and separated by a deep suture.
Sculpture consisting of numerous, elevated, slightly sigmoid

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 359

longitudinal ribs (26 on the body whorl and 20 on the penulti-
mate whorl of the holotype) with one to three low longitudinal
threads in the spaces between the ribs. Numerous fine spiral
threads are also present (about 35 on the body whorl in the
holotype) extending all over the whorls and up into the um-
bilicus. The threads on the base are a little heavier, but no
prominent cords surround the umbilicus. Aperture ovate, slightly
angular posteriorly, and in contact with the preceding whorl
only in a narrow zone. Lip thickened anteriorly and only
slightly thickened elsewhere. Umbilicus narrow and deep.
Periostracum thin, light brown. Nuclear whorls not visible.
Operculum thin, round, corneous, multispiral, light brown gen-
erally with a yellowish-green central area.

height (nuii.) width (mm.) visible whorls
holotype 3.2 » 3.0 2y2

Types. The holotype, an unique specimen, was collected alive
at R/V VEMA biology station 53 (2670 fathoms, Cape Basin,
about 300 miles southwest of Capetown, South Africa). Tt is
in the Museum of Comparative Zoology, no. 225952.

Remarks. With respect to lacking umbilical carinae, this
species is similar to B. valdiviae Thiele 1925, B. kerguelensis
Thiele 1925, B. decussata (Pelseneer) 1903, B. conica (Watson)
1886, B. erassicostata (Strebel) 1908, B. calypso (Melville and
Standen) 1912, and B. cancellata (Jeffreys) 1883. However, in
addition to being larger than any of these, the following dif-
ferences may be seen.

The spire of capensis is acute while that of erassicostata and
cancellata is obtuse. In capensis the ribs are sigmoid and the
spiral striae cross the ribs, while in valdiviae, kerguelensis, con-
ica, and decussata the ribs are straight and the spiral striae do
not cross them. B. calypso differs in having straight ribs and
two prominent lines around the penultimate whorl.

Records. Known only from the type specimen. See 'Types'.

Family CHORISTIDAE
Genus CHOEISTES Carpenter 1872

Type species: C. clegans Carpenter 1872, by monotypy.
The genus Choristes was proposed by Carpenter (in Dawson,
1872) to receive the post -Pliocene species C. elegans Carpenter.

i Upper whorl (s) broken away.

360 bulletin: museum op comparative zoology

Verrill (1882) described a living species from 255 fathoms off
Marthas Vineyard which he considered only as a variety of
elegans, viz. C. elegans var. tenera. Bush (1897), working with
additional material from off Marthas Vineyard, compared the
radula of a similar species from 390 fathoms with tenera. She
found that the "new" species possessed two single overlapping
lateral teeth (Bush, 1897, pi. 23, fig. 16) in the position occupied,
in tenera, by the single, wide, bilobed second lateral (Verrill
1882, pi. 58, fig. 27a) and that there were consequently thirteen
teeth in each transverse row in the new species instead of eleven,
the number in tenera. Bush therefore created the new genus
Choristella to receive this new species (leptalea) and another
species (brychda) represented by a single dead specimen dredged
off Marthas Vineyard in 810 fathoms and described in the same
paper.

Examination of the type specimens of the above species from
the collections of the United States National Museum has re-
sulted in the following conclusions. In shell characters tenera
and elegans are very similar except that the shell of tenem
is much thinner than that of elegans, as was pointed out by
Verrill (loc. tit.) In view of this constant and striking difference
and the fact that elegans is post-Pliocene and existed in shallow
water while tenera is recent and archibenthal, the two forms
are here considered as separate species.

On the other hand, the type specimens of leptalea and brychia,
although not equally corroded, are apparently identical to each
other and to tenera. The type localities of the three species are
all in the same general area on the continental slope south of
New England. In view of these factors, the writer considers
that the differences in the radulae which have been illustrated
(loc. tit.) may be due to different interpretations by the two
authors as to whether the second lateral was actually a single,
wide tooth with two cusps, or two separate, overlapping teeth
each with a single cusp, and that both authors probably observed
radulae of the same species.

It is, therefore, considered that the family Choristidae con-
tains only the genus Choristes, with Choristella as a synonvm.
After the addition of two species described by Dall (not dis-
cussed above) the list of species in Choristes is as follows: elegans
Carpenter 1872, tevra Verrill 1882. carp^nteri Dall 1896, pom-
pholyx (Dall) 1889, agitlhasar (sensu strieto) and agulhasae ar-
gentinae, a new species and a new subspecies to be described
below.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 361

Choristes agulhasae, new species
Plate 3, figure 1

Shell small (3.0 mm. wide), somewhat depressed, moderately
sculptured, umbilicate, semi-transparent, and very fragile.
Whorls three, convex, separated by an incised suture, and form-
ing- a flattened, obtuse spire. Color grayish white except on the
body whorl where the shell becomes transparent and longitud-
inal streaks of white appear. Sculpture consisting of fine lines
parallel to the lines of growth ; a low, centrally located, basal
carina; a second carina running up inside the umbilicus; and a
low carina on the upper part of the whorl near the suture. The
latter carina becomes obsolete on the body whorl. Aperture
large, ovate, and flattened at the inner edge. Inner lip nearly
straight, oblique, and thin. Outer lip thin and convex. Umbili-
cus rather wide and extending to the protoconch. Protoconch
small, planospiral, and consisting of about i/o whorl. Operculum
diaphanous, transparent, and apparently paucispiral.

height (mm.) width (mm.) whorls

holotype 2.0 3.0 3

Types. The holotype, an unique specimen, was collected alive
and unbroken at R/V VEMA biology station 51 (2507 fathoms,
Agulhas Basin, approximately 1000 miles southwest of Cape-
town, South Africa). It is in the Museum of Comparative
Zoology, no. 224955.

Remarks. This species, the first in this genus to be recorded
from the Southern Hemisphere, is readily distinguished from all
other species of Choristes by its rather strong basal carinae and
long, straight parietal lip. (For differential characters between
this and the following subspecies C. a. argentinae, see Remarks
under argentinae.) The other species all lack the carinae and
exhibit a curved parietal lip. In addition, pompholyx Dall is
thicker, white, and polished, while elegans Carpenter and tenera
Verrill appear to be much larger species, although, of course,
we do not know what the maximum size of agulhasae may be.

Records. The holotype is the only known specimen. See under
'Types'.

Choristes agulhasae argentinae, new subspecies
Plate 3, figures 2 and 3

Shell small (3.5 mm. wide), somewhat depressed, loosely
coiled, moderately sculptured, umbilicate, heavily eroded, and
rather fragile. Apex decollated, leaving only 1% whorls. Color

362 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

light yellowish brown except white where the outer layer of the
shell has been corroded away. Whorls convex, enlarging rapidly,
separated by a narrow incised suture, and touching only in a
narrow peripheral zone. Sculpture visible only in uncorroded
areas and consisting of fine, incremental lines parallel to the
lines of growth. There is a single, prominent, centrally located
carina on the base of the body whorl and another carina, scarcely
visible in the holotype because of corrosion, but apparently re-
volving up inside the umbilicus. Aperture large and obliquely
D-shaped. Inner lip straight, oblique and slightly thickened.
Outer lip rather thin and convex. Umbilicus rather wide, ex-
tending through the shell, and exposed apically. Protoconch
and operculum not present.

height (mm.) width (mm.) whorls

holotype 2.3 3.5 1%

Types. The holotype, without the animal and an unique
specimen, was collected from R/V VEMA biology station 12
(2805 fathoms, mid-Argentine Basin, approximately 1000 miles
east-southeast of Buenos Aires, Argentina). It is in the
Museum of Comparative Zoology, no. 224956.

Remarks. This subspecies is similar to C. agulhasae (sensu
stricto) except for the following characters. C. a. argentinae,
where it is not corroded, exhibits a somewhat thickened, semi-
translucent shell with a yellowish periostracum while in C.
agulhasae the shell is uniformly thin (much thinner than
argentinae), transparent on the body whorl, and without any
visible periostracum. In addition, the uncorroded portions of
argentinae indicate that the region of the body whorl adjacent
to the suture is slightly concave, a characteristic not seen in
agulhasae. Finally, the aperture in argentinae is more ex-
panded and the parietal lip is straighter, longer, and much
thicker than in agulhasae.

Records. Known only from the type locality. See under
'Types'.

Order MESOGASTROPODA

Family NATICIDAE
Genus AMAUROPSIS Morch 1857

Type species: Natica helicoides Johnston 1835, subsequent
designation, Dall 1909.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 363

Subgenus KerGUELENATICA Powell 1951

Type species: A. (K.) grisea (von Martens) 1878, original
designation, Powell 1951.

Amauropsis (Kerguelenatica) grisea (von Martens) 1878

Plate 1, figure 6

One dead specimen which appears to be this species was
trawled at R/V VEMA station 14 (1703 fathoms, Cape Basin,
about 400 miles northwest of Capetown, South Africa). It
was inhabited by a pale, blind, abyssal hermit crab. Another
specimen, living when collected, which unquestionably belongs
to grisea, was trawled at station 51 (2507 fathoms, Agulhas
Basin, about 1000 miles southwest of Capetown, South Africa).
This exhibited the characteristic operculum which is horny with
a prominent, thin calcareous layer over the central part, and
was otherwise identical to the published figures of grisea. A
third specimen, which is probably this species, was trawled, also
alive, at station 47 (2054 fathoms, Scotia Sea, about 60 miles
south of South Georgia). It too is very similar to the published
figures of grisea but the operculum has a heavier calcareous
layer than in typical grisea and it covers the entire outer surface.

About six other naticoid species are present in the R/V
VEMA material from stations 46, 47, 49, and 51 (1497 to 2507
fathoms, Scotia Sea and Agulhas Basin), but without type
material for comparison I do not wish to name them at this
time. They appear to belong to the group characterized by
Hedley (1916) as follows: "There is an Antarctic naticoid
group which . . . amounts to about a dozen rather featureless
species, all small, mostly uniform olive buff in color, four whorls,
a slightly raised spire, a caducous epidermis, comparatively thin,
unsculptured, except for incremental striae, without umbilical
funicle or a callus pad at the insertion of the right lip. Oper-
culum corneous paucispiral." Since the VEMA stations are in
general much deeper than any other stations in the sub-Antarc-
tic from which mollusks have been reported, identifications of
these species with previously described species would appear to
constitute large extensions of their bathymetric ranges, and such
extensions might be incorrect.

364 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Order NEOGASTROPODA

Family BUCCINULIDAE1

Genus TROMINA Dall 1918

Type species: Fusus unicarinatus Philippi, original designa-
tion.

Tromina bella abyssicola, new subspecies

Plate 2, figure 10 ; Plate 4, figure 7

Shell small (11.8 mm. long) buecinoid, moderately sculptured,
and thin. Whorls 3%, convex, separated by a rather deep suture,
and forming a spire which subtends an angle of about 70°.
Sculpturing consists of numerous, narrow, low, closely spaced
spiral ribs (about 45 on the body whorl of the holotype) and
numerous unevenly spaced axial ribs (about 75 on the body
whorl of the holotype) similar to the spiral ribs in height and
thickness, which give the surface a reticulated appearance. Aper-
ture rather large and with a wide siphonal canal, smooth and
shiny within and there exhibiting the external sculpture. Colu-
mella twisted. Parietal wall convex anteriorly and posteriorly,
concave medially with a straight central portion, and with a thin
callus over its whole surface. Outer lip thin, sharp, broadly
convex, and crenulated by the spiral ribs. Umbilicus absent.
Periostracum rather thin but prominent and yellowish brown.
Nuclear whorls 1%, forming a dome-shaped protoconch sculp-
tured with fine, slightly wavy, longitudinal ribs and nearly
imperceptible spiral lines. Operculum thin, filling about half
the aperture, light yellowish, paucispiral, and with the nucleus
sub-terminal. Radula with a tricuspid central tooth and bi-
cuspid marginals (PI. 4, fig. 7).

height (mm.) width (mm.) whorls

holotype, station 51 11.8 7.5 3:'^

para type, station 51 7.2 4.9 3%

Types. The holotype and one paratype, the only specimens
known, were collected alive at R/V VEMA biology station 51
(2507 fathoms, Agulhas Basin, about 1000 miles southwest of
Capetown, South Africa). The holotype is in the Museum of
Comparative Zoology, no. 224954. The paratype is in the Na-
tional Museum of Canada, no. 4739.

i The writer is following Powell (1951) in the use of this family name.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 365

Remarks. T. bella abyssicola is similar to T. bella Powell
1951 (82-152 fathoms, from four localities near the Falkland
Islands) in most of its characters and is apparently closely
related to it. The observed differences seem to be entirely suf-
ficient to justify its subspecific status however, especially in
view of the great divergence in depth and the great distance
between the localities.

T. b. abyssicola is a much thinner and more delicate species
than T. bella. The external sculpturing shows clearly inside
the aperture in abyssicola but does not show in bella. The
parietal wall in abyssicola has a straight portion in the center
while in bella the central region is rather evenly concave. In
addition, the central tooth of the radula of abyssicola bears a
larger central and two markedly smaller lateral cusps and the
two cusps of each marginal are of approximately equal size
(PI. 4, fig. 7). In bella however, the central tooth bears three
cusps of the same size and the inner cusp of the marginals is
larger than the outer (Powell 1951, p. 194, fig. 72). The shape
of the teeth is also somewhat different in the two subspecies.

The shell of this subspecies is very similar to some species in
the taenioglossate archibenthal and abyssal genus Oocorys
(family Tonnidae, see Turner, 1948) and represents another
case of parallel evolution among unrelated groups. Certain of
the published records of Oocorys from southern high latitudes
may well be based on species of Tromina.

Records. Known only from the type locality. See under
'Types'.

Tromina traverseensis, new species
Plate 2, figure 8

Shell small (6.6 mm. long), buccinoid, weakly sculptured,
and thin. Whorls 3%, convex, shouldered, separated by a deep
suture and forming a spire which subtends an angle of about
80°. Sculpture consisting of many narrow, low, rounded spiral
ribs (about 43 on the body whorl of the holotype) which appear
as whitish lines on a pale buff background, crossed by very
fine, crowded lines of growth. Aperture large (about % the
length of the shell), with numerous narrow, shallow grooves
within, resulting from the external ribbing, and w7ith a wide
siphonal canal. Columella twisted. Parietal wall convex an-
teriorly and posteriorly and concave centrally and glazed with
a thin callus which does not obscure the sculpturing beneath.

height (mm.)

width (mm.)

holotype, station 49

6.6

4.5

paratype, station 49

2.7

2.0

366 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Outer lip thin, sharp, broadly convex and lightly cremilated by
the spiral sculpture. Umbilicus small and nearly closed by the
flexion of the parietal callus. Periostracum thin, yellowish
brown, and occurring mainly between the spiral ribs. Nuclear
whorls V/2, nearly smooth, and forming a dome shaped proto-
conch. The holotype, containing the soft parts, has no oper-
culum.

whorls
3%
2%

Types. The holotype and paratype, respectively with and
without the soft parts, were collected at R/V VEMA biology
station 49 (1497 fathoms, south of Traverse Island, South
Sandwich Islands). The holotype is in the Museum of Com-
parative Zoology, no. 224963. The paratype is in the National
Museum of Canada, no. 4738.

Remarks. T. traverseensis is somewhat similar to T. simplex
Powell 1951 in sculpturing, but traverseensis has strongly con-
vex, shouldered whorls, an umbilicus, is very thin, and shows
the external sculpturing within the aperture, while simplex has
only lightly convex, unshouldered whorls, lacks an umbilicus,
is substantially thicker, and does not exhibit the internal sculp-
turing within the aperture. Compared with T. bella abyssicola
Clarke, traverseensis has much finer sculpturing and possesses
an umbilicus while abyssicola does not.

Records. The only known specimens are the holotype and
paratype. See under 'Types'.

Shells of what now appear to be three additional new species
of Tromina (or possibly Notoficula Thiele 1917) are in the
R/V VEMA material. Each is represented by a single specimen,
of which two are somewhat broken empty shells and probably
immature (from VEMA stations 47 and 53) and one is very
young and contains the soft parts (station 47). So little is
known of the degree of variation to be expected among these
Antarctic whelks that it is quite uncertain whether these will
prove to be outside the range of variation of known species or
not. If they are similar to northern whelks in this regard
(Bnccinum, Neptunea, Coins, etc.)., much variation may be
expected. For this reason, I do not wish to describe them until
more material is available and in further discussions will refer
to them simply as Tromina a, b, and c, respectively.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 367

Class PELECYPODA

Order PROTOBRANCHIATA

Family NUCULIDAE

Genus NUCULA Lamarck 1799

Type species : Area nucleus Linne, by monotypy.

Nucula turnerae, new species

Plate 2, figures 2 and 3

Shell minute (1.8 mm. long), ovate, sub-inflated, nearly
smooth, thin and with a broadly curved hinge plate bearing few
teeth. Valves fragile and semi-transparent. Outline ovate ex-
cept for the protruding umbones (which are placed about 36 per
cent of the length from the posterior end), a slightly pointed
anterior end (the longer end), and a somewhat flattened posterior
end. Sculpture consisting of numerous fine lines of growth and
microscopic radial lines. Larval valves very small, caplike and
covering only the dorsal part of the umbones. Periostracum
very thin and pale brown. Inner surface shiny, exhibiting
well-marked muscle scars and a simple pallial line. Shell margin
smooth and covered by periostracum. Hinge plate broadly
curved, narrowed anterior to the chondrophore. Anterior row
with three and posterior row with two, medium-sized, pyramidal,
dorso-ventrally compressed taxodont teeth. Hinge plate ex-
panded at the chondrophore which is small, triangular-ovate,
and directed obliquely anteriorly. Umbones excavated.

length (mm.) height (mm.) width (mm.)

holotype, station 12 1.8 1.5 1.0

Types. The holotype, an unique specimen, was collected alive
at R/V VEMA biology station 12 (2805 fathoms, Argentine
Basin, about 1000 miles east-southeast of Buenos Aires, Argen-
tina). It is in the Museum of Comparative Zoology, no. 224959.

Remarks. The simple, plain and fragile shell of turnerae
renders it distinct from all other abyssal nuculas from the
South Atlantic Ocean. In general appearance it approaches
N. pernambucensis Smith 1885 from off Recife, Brazil, in 675
fathoms (CHALLENGER, sta. 120), but that species is larger,
much more quadrate and bears about eleven teeth in each valve
while turnerae bears only five. It is possible that turnerae is
immature, but the thin, fragile, nearly unsculptured shell is

368 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

typical of deep abyssal species and the tiny attached larval
shell suggests that the holotype is probably adult or nearly so.

This species is named in honor of Dr. Ruth D. Turner who
has generously aided the writer in numerous matters connected
with this work.

Records. The holotype is the only known specimen. See under
'Types'.

Genus PrONITCULA Hedley 1902

Type species: Pronucula decorosa Hedley 1912, original desig-
nation.

Pronucula benguelana, new species
Plate 3, figures 9 and 11

Shell nuculiform, slightly inflated, small (about 3.5 mm.
long), radially sculptured, thin and with a curved hinge plate.
Valves semi-transparent and showing the teeth, intestine, etc.
through the shell. Outline triangular-ovate, rounded and
slightly pointed posteriorly, broadly curved ventrally, more
abruptly curved anteriorly (sometimes slightly pointed) and
with prominent, inflated umbones placed about 40 per cent of
the length from the posterior end. Adult sculpture consists of
numerous (about 65 to 75) narrow, rounded radial ribs over
the whole central area crossed by crowded, microscopic lines of
growth. Larval shells white, persistent, large (about 30 per
cent of the height of tbe adult) and sculptured with fine con-
centric lines and microscopic radial lines. Periostracum thin,
lustrous and light yellowish brown. Inner surface nacreous
and exhibiting the external radial ribs. Muscle scars and simple
pallial line well marked. Shell margin smooth, not crenulated.
Hinge plate nearly straight posteriorly, rather sharply curved
centrally and broadly curved anteriorly. Posterior row with
six and anterior row with nine columnar, pointed, slightly
curved teeth. Chondrophore small, triangular, approximately
vertical and close to tbe innermost taxodont teeth. Umbones
deeply excavated.

length (mm.) height (mm.) width (mm.)
holotype, station 14 3.8 3.4 2.0

pavatype, station 14 3.3 3.0 1.8

paratype, station 14 3.2 2.9 1.8

paratype, station 14 2.9 2.4 1.6

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 369

Types. The holotype and 26 paratypes, all living specimens,
plus a single valve, were collected at R/V VEMA station 14
(1703 fathoms, Cape Basin, about 400 miles northwest of Cape-
town, South Africa). The holotype is in the Museum of Com-
parative Zoology, no. 224964. Paratypes are in the Museum of
Comparative Zoology and the National Museum of Canada.

Remarks. Pronucula benguclana is one of a widespread deep
sea complex which includes P. notobentlialis (Thiele) 1912 from
1490 fathoms from the western end of the Eastern Indian
Antarctic Basin and P. profunclorum (Smith) 1885 from 2050
fathoms from the mid-North Pacific. It differs from profun-
clorum in that profandorum is much more pointed posteriorly
and the beaks are more centrally located. Conversely, from
notobentlialis it differs in that the posterior extremity is much
more rounded and the beaks are located more anteriorly in
that species; also notobenthalis is much more oblique.

The name benguclana refers to the type locality, which is
beneath the region traversed by the Benguela Current.

Records. In addition to the holotype and paratypes, there are
thirteen specimens from station 47 (2054 fathoms, Scotia Sea,
about 60 miles south of South Georgia) which are doubtfully
referred to this species. They are very similar to the specimens
from the type locality except that the larval shells are propor-
tionately larger and the adult outline is more broadly curved
posteriorly. Such differences may be expected between isolated
populations of abyssal mollusks because of very restricted gene
flow between them. In this case the differences are relatively
minor and I prefer not to designate this population as a distinct
subspecies.

Family MALLETIIDAE

Genus MALLETIA Desmoulins 1832

Type species : Malletia chilensis Desmoulins, by monotypy.

Malletia pallida Smith 1885

This species was taken at R/V VEMA biology station 47
and 52 (2054 fathoms, Scotia Sea, approximately 60 miles
south of South Georgia; and 2711 fathoms, Agulhas Basin, ap-
proximately 800 miles southwest of Capetown, South Africa, re-
spectively). A series of 15 specimens was collected at station
52, with individuals ranging from 3.0 to 19.5 mm. in length.

370 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

The larger specimens are identical to the figures in Smith
(1885, pi. 20, figs. 8, 8a). The smaller specimens are very simi-
lar to the three small specimens from station 47, which otherwise
would not have been recognized as belonging to pallida. The
type locality is CHALLENGER station 137, 2550 fathoms,
35°59' S., 1°34' E., in the Cape Basin, not far from VEMA
station 52.

Malletia johnsoni, new species
Plate 2, figure 6

Shell small (about 6 mm. long), rectangularly ovate, com-
pressed, nearly smooth, very thin and fragile, and with a nar-
row, weak and almost straight hinge plate. Valves somewhat
translucent, thin and brittle. Dorsal margin nearly straight;
anterior margin rounded and curving smoothly to the ventral
margin which is less sharply rounded; posterior broader than
anterior and truncated. Umbones rather small, placed a little in
front of center. Sculpture consisting only of fine lines of
growth. Periostracum thin, shiny, and pale yellowish brown.
Ligament extending from slightly in front of the beaks to near
the posterior margin. Inner surface white, only slightly lustrous
and with muscle scars and a relatively large pallial sinus located
above the midline of the valves. Shell margin finely crenulated.
Hinge plate narrow, nearly straight, a little bent down an-
teriorly, wider distally than centrally, and bearing about 7
anterior and 10 posterior small V-shaped taxodont teeth. Teeth
larger distally and becoming obsolete and vanishing near the
beaks. Umbones not excavated.

holotype, station 22
paratype, station 22
paratype, station 22

Types. The holotype and six paratypes were collected alive
at R/V VEMA biology station 22 (1675 fathoms, Angola Basin,
about 175 miles west of Banana, Belgian Congo). The holotype
is in the Museum of Comparative Zoology, no. 224961. Para-
types are in the National Museum of Canada, no. 4737.

Remarks. Malletia johnsoni is characterized by its small size,
its compressed, thin, and fragile valves, and its broad, truncate
posterior end. It resembles the North Atlantic M. ootusata
Sars in general, but that species is not broader posteriorly, the

:h (ruiu.)
4.7

height (ram.)
3.1

width (mm.)
1.4

6.2

3.8

1.6

4.0

2.5

1.2

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 371

beaks are farther forward and the valves are not convex. M.
dilatata Philippi is much more heavily sculptured and the pos-
terior end is more sharply truncated. M. johnsoni is not similar
to any South Atlantic species.

This species is named in honor of Richard I. Johnson, who
for many years has been a most enthusiastic and careful worker
in malacology.

Records. In addition to the type lot, two specimens which
probably belong to this species were collected at station 54 (993
fathoms, Cape Basin, about 65 miles northwest of Capetown,
South Africa). They are very similar to the types except that
the valves are a little larger (7.0 mm. long) and significantly
more transparent. Seen through the shell, the number of teeth
also appears to be a little greater, but this is no doubt because
of the greater length.

Malletia concentrica Thiele 1912

Ten specimens of one species of Malletia ranging in length
from 2.0 to 5.3 mm. were taken at R/V VEMA biology station
49 (1497 fathoms, Atlantic Indian Antarctic Basin, south of
Traverse Island, South Sandwich Islands) and three other
specimens ranging from 1.5 to 2.7 mm. in length were taken at
station 16 (1660 fathoms, Cape Basin, approximately 175 miles
west-northwest of Luderitz, South-West Africa). The smaller
specimens resemble Thiele 's figures (1912: pi. 17, figs. 24, 24a)
of concentrica very closely (the holotype is 2.75 mm. long) and
probably belong to that species. The type locality of concentrica
is 1872 fathoms northeast of Gauss Station in the western end
of the Eastern Indian Antarctic Basin. Larger specimens of
concentrica from station 49 are expanded post-basally and are
quite unsymmetrical in shape. The species is markedly convex,
the largest specimen measuring 5.3 mm. in length, 3.8 mm. in
height and 2.8 mm. in width.

Genus TiNDARIA Bellardi 1875
Type species: Tindaria arata Bellardi, by monotypy.
Tindaria virens Dall 1890

At R/V VEMA biology station 22 (1675 fathoms, Congo
Canyon, Angola Basin, about 175 miles west of Banana, Belgian
Congo) twelve living specimens were collected which agree
closely with paratypes of Tindaria virens Dall. The only

372 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

observable difference is that the hinge teeth are a little heavier
in the VEMA specimens than in virens, but this difference is
probably too slight to be given taxonomic significance. T. virens
was taken by Dall off southern Chile in 122 to 449 fathoms.

Tindaria championi, new species
Plate 2, figures 1 and 4

Shell small (5.8 mm. long), veneriform, sub-inflated, moder-
ately strong, rather weakly sculptured and with an angular
hinge plate bearing taxodont hinge teeth. Outline oval except
for the prominent beaks which are placed about 40 per cent of
the way from the anterior end. Sculpture consisting of numer-
ous narrow, very low, rounded concentric ribs, fine near the
umbones and becoming coarser near the ventral margin, and
crowded, exceedingly fine radial lines. Periostracum thin, light
brown and beautifully iridescent. Ligament narrow, about 18
per cent the length of the shell, and extending posteriorly from
under the umbones. Inner surface white, sub-nacreous, with
prominent muscle scars, pallial line and pallial sinus. Shell
margin smooth. Hinge plate strong, bent and a little narrowed
under the umbones and bearing seven anterior and nine pos-
terior, erect, V-shaped, conical taxodont teeth. Teeth much
higher in the center of each row, becoming obsolete near the
umbones, and not meeting under the umbones. Umbones ex-
cavated.

length (nun.) height (mm.) width (mm.)
holotype, station 12 5.8 4.8 2.8

Types. The holotype was taken alive at R/V VEMA biology
station 12 (2805 fathoms, Argentine Basin, about 1000 miles east-
southeast of Buenos Aires, Argentina). It is in the Museum of
Comparative Zoology, no. 224957.

Remarks. Tindaria championi is another species of the veneri-
form, concentrically sculptured group in Tindaria which has a
prominent pallial sinus (see T. creous Clarke 1959b). T. cham-
pioni may be distinguished from T. ercbiis by the much heavier
sculpture, many more teeth, regularly curved central portion of
the hinge plate, and larger size of erchns. It differs from T.
antarctica Thiele and Jaeckel in shape (subrectangular in an-
tarctica and oval in championi) , in the more anterior placement
of the umbones in antarctica, and the number of posterior teeth
which is much greater in antarctica. It is not closely similar to
any other South Atlantic species.

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 373

This species is named for Merrill E. Champion who for many
years has given much time and energy to the study of Recent
mollusks at the Museum of Comparative Zoology.

Records. In addition to the holotype from station 12, seven
additional specimens were collected at station 14 (1703 fathoms,
Cape Basin, about 14 miles northwest of Capetown, South
Africa). They differ from the holotype in their smaller size
and less iridescent periostracum but are identical in all other
respects.

Tindaria Antarctica Thiele and Jaeckel, 1931

One living specimen of this species was collected at R/V
VEMA biology station 47 (2054 fathoms, western end of the
Atlantic Indian Antarctic Basin, about 60 miles south of South
Georgia) and another living specimen and two extra valves
were collected at station 48 (1902 fathoms, about 100 miles
southeast of South Georgia). They agree perfectly with the
description and figure in Thiele and Jaeckel (1931, pi. 8, fig. 71)
except for a slight difference in the number of teeth (8 anterior
and 18 posterior instead of 6 and 20 as in (infarct tea) and their
somewhat smaller size (8.5 mm. long instead of 11.5 mm.). The
type locality of antarctica is VALDIVIA station 152 (2535
fathoms at 63°16.5' S., 57°51' E., eastern end of the Atlantic
Indian Antarctic Basin).

CLENCHARIA, new subgenus

Type species: Tindaria (Clcncharia) diaphana Clarke.

Shells small, ovate, thin, transparent and with numerous flat-
tened taxodont teeth. Pallial sinus well developed. Taxodont
teeth wedge shaped and flattened laterally. Sculpture concentric
and microscopic.

Clcncharia differs from Tindaria (sensu stricto) principally in
the form of the teeth, the sculpturing and the thickness and
transparency of the shell. The teeth in Clencharia are flattened
and rectangular in cross-section while those of Tindaria are
V-shaped in cross-section. The sculpturing in Clencharia is
microscopic ; in Tindaria it is not microscopic and is often prom-
inent. The shell of Clcncharia is very thin, diaphanous and
transparent while that of Tindaria is relatively thick and not
transparent.

This subgenus is named in honor of Dr. William J. Clench, a
fine teacher and an outstanding authority on mollusks with
whom the author has had the privilege of being associated.

374 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Tindaria (Clencharia) diaphana, new species
Plate 1, figure 2

Shell small (5.6 nrni. long), sub-ovate, inflated, nearly smooth,
very thin, transparent and with a weak hinge plate bearing
about 15 flattened chisel-shaped teeth. Valves completely trans-
parent and clearly exhibiting the entire animal within. Outline
sub-ovate, broadly rounded postero-ventrally and ventrally.
gradually rounded and straighter dorsally and more sharply
curved anteriorly and postero-dorsally. Umbones rounded, in-
flated, and projecting above the hinge line at a point about
30 per cent of the distance from anterior to posterior. Sculpture
consisting only of microscopic concentric lines and ridges of
growth. Periostracum very thin, transparent and iridescent.
Ligament very thin, extending posteriorly from under the um-
bones. Larval valves tiny and white. Interior smooth except
for microscopic growth lines, glossy, and exhibiting well marked
muscle scars, pallial line and pallial sinus. Shell margin smooth.
Hinge plate compressed and narrow except a little wider an-
teriorly and somewhat thickened and sigmoid under the beaks,
but otherwise following the dorsal margin of the valves. Hinge
plate bearing six anterior and nine posterior dorso-ventrally
compressed, chisel-shaped taxodont teeth. Teeth well developed
but rather small (largest near the center of each row) and
becoming obsolete and not meeting under the beaks. Umbones
excavated.

holotype, station 52

length (mm.)
5.6

height (mm.)
4.1

width (mm.)
2.8

paratype, station 52

5.7

4.2

2.9

paratype, station 52

4.4

3.2

1.9

Types. The holotype and two paratypes were taken at R/Y
VEMA biology station 52 (2711 fathoms, Agulhas Basin, about
800 miles southwest of Capetown, South Africa). The holotype
(with a flake of orange paint from the dredge inside the shell)
is in the Museum of Comparative Zoology, no. 224965. The
paratypes are in the National Museum of Canada, no. 4741.

Remarks. Tindaria (Clencharia) diaphana is characterized
by its frail, transparent, bulbous shell and by its peculiar chisel-
shaped teeth. It can be differentiated from all other fragile
taxodonts (e.g. Glomus, Phaseolus, Sarepta, etc.) by the general
shape of the shell and the form of the teeth. For differences
between this and Tindaria (sensu stricto), see under Clencharia.

Records. Known only from station 52. See under 'Tj^pes'.

CLARKE : ABYSSAL SOUTH ATLANTIC MOLLUSKS 375

Family NUCULANIDAE
Genus NUCULANA Link 1807

Type species: Area rostrata Chemnitz (= Leda pernula Miil-
ler), by monotypy.

Nuculana ultima Smith 1885

Nuculana ultima Smith 1885, Challenger Report, LameUibranchs, p. 324,

text fig. (2740 fms., southwest of Canary Islands).
Leda (Ledella) modesta Thiele and Jaeckel 1931, Muscheln der deutschen

Tief see-Expedition, p. 202, pi. 2, fig. 30 (1245 fms., Gulf of Guinea).
Leda (Ledella) spreta Thiele and Jaeckel 1931, loc. cit., p. 203, pi. 2, fig.
32 (69 fathoms, Agulhas Bank).
Living specimens of Nuculana ultima were taken by the
VEMA at stations 12, 23, 47, 51, and 52 (2504 to 2805 fathoms;
Argentine, Angola and Agulhas basins and Scotia Sea). Sev-
eral specimens were collected at each locality and large series
were obtained at stations 12 and 52. The species is quite variable
as to the presence or absence of a short rostrum, in the strength
of the hinge plate, and especially in the thickness and inflation
of the shell. Apparently when the species reaches adult length
and height further growth takes place at the inner edge of the
shell and it becomes progressively more obese. At the same
time the whole shell thickens. In one lot (station 52) this has
resulted in sufficient variation so that width/height ratios vary
from .65 to .95 among adult specimens. Nuculana ultima Smith
was described from a single specimen, and modesta and spreta
Thiele and Jaeckel were also represented by only a few shells.
These authors apparently obtained isolated examples of an
unusually variable species and under such conditions they ap-
peared distinct.

Subgenus THESTYLEDA Iredale 1929
Type species: Leda ramsayi Smith, original designation.
Nuculana (Thestyleda) louiseae, new species
Plate 1, figure 7

Shell medium sized (19 mm.), rather thin, narrow, rostrate,
compressed, strongly sculptured and with numerous oblique,
lamellar, taxodont teeth. Valves ovate anteriorly, with a long,
arcuate, posterior rostrum and with small umbones placed close
together and about 30 per cent of the length from the anterior

376 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

end. Rostrum narrow, truncate and bilobed distally, concave
dorsally and ventrally and surmounted with two prominent,
curved carinae on each valve running from the umbones to the
distal end. The lower carina is larger and becomes double as
it proceeds posteriorly. Carinae crossed by crisp, narrow, wave-
like ridges, which on the upper carina are convex and on the
lower carina are concave dorsally and sharply convex ventrally.
Ridges continuing on the disc as narrow, rounded, closely spaced
ribs following lines of growth. Between the long, thin, external
ligament and the rostral carinae is a smooth, almost unsculp-
tured area, extending from the umbo and becoming narrow and
obsolete near the posterior end. Periostracum thin, light grayish
yellow. Inner surface of valves mostly white, nacreous, showing
the external sculpturing and numerous, weak radial lines. An-
terior muscle scar circular and faint, posterior scar irregularly
ovate, and pallia! line indistinct. Hinge teeth compressed,
v-shaped, lamellar, very oblique, rather high and firmly inter-
locking, about 12 anteriorly and 20 posteriorly. Internal
resilium in two separate but adjacent parts, the anterior part
small and round, the posterior part large and saddle shaped.
Umbones not excavated.

length (mrn.) height (mm.) width (mm.)
holotype, station 12 19.0 7.0 3.5

Types. The holotype, an unique specimen, was collected at
R/V VEMA biology station 12 (2805 fathoms, Argentine Basin,
about 1000 miles east -southeast of Buenos Aires, Argentina).
It is in the Museum of Comparative Zoology, no. 224958.

Remarks. Nuculana louiseae is characterized by its very long
and narrow rostrum and its peculiar sculpture. Apparently
it is not closely related to any other living species. It is some-
what similar to Leda longicaudaia Thiele 1912 from the western
end of the Eastern Indian Antarctic Basin, but in that species
the rostrum is less extended, differently sculptured, and convex
ventrally, while in louiseae it is concave. Compared with
Nuculana scalata Prashad, from 500 fathoms near the Lesser
Sunda Islands, northeast Indian Ocean, louiseae is higher and
much more expanded anteriorly and with the umbones placed
more posteriorly.

This species is named in honor of my wife, Louise R. Clarke,
who has been a constant source of help in my work on the
Mollusca.

Records. Known only from the type locality. See under
'Types'.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 377

Genus SPINULA Dall 1908

Type species: Leda (Spinula) calcar Dall, original designa-
tion.

Spinula subexcisa (Dautzenberg and Fischer) 1897

Specimens which are here referred to Spinula subexcisa were
collected at R/V VEMA biology stations 12, 14, 18, 19, 51 and
52 (1510-2805 fathoms, Argentine, Agulhas and Cape basins
and Walvis Ridge) .

Jeffreys (1876, 1879) recorded Malletia excisa (Philippi) in
his reports on the VALOROUS and the LIGHTNING and
PORCUPINE expeditions from 1125 to 1785 fathoms in the
West Europe and Canaries basins. Comparisons with the
Jeffreys Collection now in the United States National Museum
show that the VEMA specimens cited here are identical to
M. excisa, as Jeffreys understood it. Philippi 's figure (1844,
pi. 15, fig. 4) is of a different species however; it is a Tertiary
fossil which is much more deeply excavated below the rostrum
than is the VEMA species. This lack of agreement between the
fossil and the recent abyssal species was recognized by Dautzen-
berg and Fischer (1897: 203) who proposed "var. subexcisa"
for the modern species previously recorded by Jeffreys and
others as excisa.

I consider that the present species is so different from the
fossil excisa that it certainly deserves specific rank.

Spinula messanensis ("Seguenza" Jeffreys) 1879

A single specimen was collected alive at R/V VEMA biology
station 20 (2707 fathoms, Angola Basin, about 675 miles west
of Walvis Bay, South-West Africa). It is very close to
messanensis which has been recorded from the West Europe and
Canaries Basin in 276 to 1731 fathoms (Locard 1897) and
although it may later prove to be significantly (and subspecific-
ally) different when more material is obtained this question
cannot be decided now.

Order ANISOMYARIA
Fairiily MYTILIDAE
Genus DaCRYDIUM Torell 1859
Type species : Madiola vitrea Moller, by monotypy.

378 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Dacrydium albidum Pelseneer 1903

Living specimens of what appear to be this species were col-
lected at R/V VEMA stations 16, 18 and 25 (Cape, Angola
and Guinea basins, 1660 to 2315 fathoms). They are almost
completely transparent and reveal the entire animal within.
In this they differ from albidum, and in a group so lacking in
taxonomic characters this may be significant, but for the present
they are considered as abyssal and depauperate representatives
of that species.

Family PECTINIDAE

Genus PSEUDAMUSIUM Morch 1853

Type species: Ostrea hybrida Gmelin, subsequent designa-
tion, Dall 1898.

Pseudamusium pteriola (Melville and Standen) 1907

A single valve of this species was collected at R/V VEMA
biology station 46 (2030 fathoms, Scotia Sea, about 60 miles
south of South Georgia). The type locality of pteriola is:
Scotia Bay, South Orkney Islands, 9 to 10y2 fathoms. Ap-
parently this valve is advectitious.

Genus PROPEAMUSSIUM de Gregorio 1883

Type species: Pecten (Propeamussium) cecilae de Gregorio,
by monotypy.

Propeamussium (Parvamussium) octodecimliratum
(Melville and Standen) 1907

One living specimen plus a fragment of what appears to be
this species were taken at R/V VEMA biology station 51 (2507
fathoms, Agulhas Basin, about 1000 miles southwest of Cape-
town, South Africa). It differs from octodecimliratum in the
possession of 13 instead of 18 internal ribs, but since it is only
4.3 mm. high (the holotype of octodecimliratum is 9.0 mm.
high) this difference is probably attributable to age. Occa-
sional specimens of Propeamussium (e.g. P. dalli Smith) inter-
polate additional ribs with growth, and this may occur in the
present species. The type locality of P. octodecimliratum is :
2500 fathoms, 67°33' S., 36°35'W., western end of Atlantic
Indian Antarctic Basin.

CLARKE: ABYSSAL SOUTH ATLANTIC MOLLUSKS 379

Family LIMIDAE

Genus LlMATTJLA S. Wood 1839

Type species: Pecten subauriculata Montagu, subsequent
designation, Gray 1847.

Limatula simillima (Thiele) 1912

One valve of this species occurred at R/V VEMA biology sta-
tion 14 (1703 fathoms, Cape Basin, about 400 miles northwest
of Capetown, South Africa). It was perforated with a neat,
round hole, apparently caused by a gastropod. Such perfora-
tions are rare among abyssal mollusks and this specimen is
probably advectitious in the deep sea. The type locality is Gauss
Station in the southwestern part of the Eastern Indian Ant-
arctic Basin.

Order EULAMELLIBRANCHIA
Family THYASIRIDAE

Genus THYASIEA "Leach" Lamarck 1818
Type species : Tellina flexuosa Montagu, by monotypy.
Thyasiba febbuginosa (Forbes) 1844

Living specimens were collected at VEMA stations 16, 22 and
47 (1660, 1675 and 2056 fathoms in the Cape and Angola basins
and the Scotia Sea). They were particularly common at station
16. They closely resemble specimens of the wide-ranging North
Atlantic archibenthal and abyssal species T. ferruginosa, and
are tentatively referred to that species.

Order SEPTIBRANCHIATA
Family POROMYACIDAE

Genus POEOMYA Forbes 1844

Type species: P. anatinoidea Forbes (= P. granulata Nyst),
by monotypy.

Poeomya sublevis Verrill 1884

A single living specimen of Poromya which is apparently
identical with Dall's holotype of P. microdonta (= P. sublevis
Verrill) was collected at R/V VEMA biology station 25 (2315
fathoms, Guinea Basin, approximately 675 miles south of

380 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Accra, Gold Coast). The type locality is in the North American
Basin, 125 miles off Chesapeake Bay, Virginia, in 1685 fathoms.
Dall (1889b) has extended the range of this species to "Pata-
gonia" and "West America" but without station data or other
explanation. These records need confirmation.

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1944. Prosobranchia. 1639 pages. Berlin, Gebriider Borntraeger.
White, K. N.

1942. The anatomy of Cyclo.strema bushi D. and F. Proc. Malacol.
Soc. London 25: 89-94.

WOODRING, W. P.

1928. Miocene mollusks from Bowden, Jamaica. Parts 1 and 2. Car-
negie Inst. Washington, Publ. 366, 385, 786 pp., 68 pis.
Yonge, C. M.

1939. The protobranchiate Mollusca; a functional interpretation of
their structure and evolution. Philos. Trans. Roy. Soc. London,
no. 566,230: 79-147.

PLATES

Plate 1

Figure Page

1. Broolcula (Vetulonia) galapagana Dall 354

Holotype, about 15 X.

2. Tindaria (Clencharia) diaphana, n. sp 374

Holotype, about 8.6 X.

3. Puncturella (Fissurisepta) agulhasae, n. sp 347

Holotype, basal view, about 7.3 X.

4. Broolcula (Benthobroolcida) capensis, n. sp. 358

Holotype, about 14 X.

5. Broolcula (Broolcula) stibarochUa Iredale 354

Holotype, about 35 X.
(i. Amauropsis (Kcrguelcnatica) grisea (von Martens) 303

E/V VEMA Station 14, about 3.5 X.
7. Xuculana (Thestyleda) louiseae, n. sp 375

Holotype, about 5.1 X.

Plate 1

Plate 2

Figure Page

1. 4. Tindaria championi, n. sp 372

1. Holotype, right valve, about 6.7 X.

4. Holotype, right valve, about 6.7 X.

2. ?>. Nucula turnerae, n. sp 367

2. Holotype, left valve, about 21 X.

3. Holotype, right valve, about 21 X.

5. 7. Seguensia elegans Jeffreys ... 350

5. M/V THETA Station 9 (2843 fms., 185 miles west of
Bermuda), about 7.1 X.

7. R/V VEMA Station 12, about 6.6 X.

6. Mdlletia johnsoni, n. sp. 370

Holotype, about 9.2 X.

8. Tromina traverseensis, n. sp 365

Holotype, about 9.1 X.

9. Puncturella (Fissurisepta) agulhasae, n. sp. 347

Holotype, about 6.4 X.

1(1. Tromina bella abyssicola, n. ssp 364

Holotype, about 5.9 X.

ityas^

\ 7

Mb.

: i " . ■ i.

Plate 2

Plate 3

Figure Page

1. Choristes agulhasae, n. sp. 361

Holotype, about 10 X.

2. 3. Choristes agulhasae argenlinae, n. ssp. 3(>1

2. Holotype, about 11 X.

3. Holotype, about 9 X.

4. Abyssogyra vemae, n. sp. 353

Holotype, about 18 X.

5. Seguenzia elegans Jeffreys 350

R/V VBMA Station 23, about 9.5 X. Specific identity not
certain.

6. Seguenzia eritima Verrill 350

R/V VEMA Station 18, about -45 X.

7. BrooTcula (Bonthobrookula ) powelli, n. sp. 355

Holotype, basal view, about 18 X.

8. BrooTcula (BenthobrooTc ula) exquisita, n. sp. 35G

Holotype, basal view, about 18 X.

9.11. I'ronueula benguelana, n. sp. ... 3G8

9. Holotype, right valve, about 10 X.
11. Holotype, left valve, about 10 X.

10. Seguenzia antarctica Thiele ... 350

K/V VEMA Station 18, about 28 X.

Plate 3

Plate 4

Figure Page

1. Brookula (Benthobrookula) powelli, n. sp. 35")

Paratype, apical view, about 20 X.

2. Brookula (Benthobrookula ) exquisita, n. sp 356

Holotype, about 20 X.

3. Brookula (Benthobrookula ) larnonti, n. sp 357

Holotype, about 20 X.

4. Scffitenzia louiseae, n. sp. 351

Holotype, about 18 X.

5. 8. Cocculina superba Clarke, 1960. R/V VEMA Station 212

3334 fathoms, Argentine Basin.

5. Holotype, basal view, about 1.25 X.

8. Holotype, apical view, about 1.25 X.
(I. Segucnsia carinata Jeffreys 350

R/V VEMA Station 14, about 15 X.
7. Raclula of Tromina bella abyssicola, n. ssp. 364

Radula from holotype.
9. Raclula of BrooTcula (Benthobrookula) powelli, n. sp. 355

Radula- from paratype, R/V VEMA Station 51.

Plate i

Bulletin of the Museum of Comparative Zoology

AT HAEVAED COLLEGE

Vol. 125, No. 13

THE GENUS MICRATHENA (ARANEAE, ARGIOPIDAE)

IN CENTRAL AMERICA

By Arthur M. Chickering

Albion College, Albion, Michigan

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

November, 1961

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WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

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1945 — Vol. 2, no. 26 is current.

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and future volumes will be published under Museum auspices.

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is available.

Bulletin of the Museum of Comparative Zoology

AT HAEVARD COLLEGE

Vol. 125, No. 13

THE GENUS MIC R ATHENA (ARANEAE, ARGIOPIDAE)

IN CENTRAL AMERICA

By Arthur M. Chickerin.g

Albion College, Albion, Michigan

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

November, 1961

No. 13 — The Genus Micrathena (Araneae, Argiopidae)
in Central America

By Arthur M. Chickering

The genus Micrathena Sundevall, 1833 with its numerous
species, more or less conspicuous webbing, and often gaudy color-
ation constitutes an important segment of the arthropod fauna
of Central America. For more than thirty years, during repeated
visits to Panama and nearby regions, I have been collecting mem-
bers of this genus with the intention of publishing the results of
my study of all species appearing in my collections. Upon the
advice of other araneologists I have recently extended my study
to include the whole of Central America. In order to aid me
in this work all specimens of the genus in the Museum of Com-
parative Zoology at Harvard College have been turned over to
me for study. During the summer of 1958, while on a fellowship
from the John Simon Guggenheim Memorial Foundation, I was
able to work on the extensive collections in the British Museum
(Natural History) where I had access to most of the material
which furnished the basis for the work of the two Pickard-Cam-
bridges published in the Biologia Centrali-Americana. A few
days were also spent in the Oxford University Museum where a
portion of the Pickard-Cambridge collection is kept.

Acknowledgements are due and gratefully rendered to the fol-
lowing named individuals and organizations for their numerous
courtesies, support, and encouragement of my work for many
years: The John Simon Guggenheim Memorial Foundation for
its financial aid during two successive years ; Dr. G. Owen Evans,
Department of Zoology, British Museum (Natural History),
and his staff of co-workers all of whom gave me very friendly
encouragement and the loan of important specimens for the
pursuit of my work; Professor G. C. Varley and his staff of
co-workers at the Oxford University Museum who gave me access
to valuable specimens for a brief period and then extended ma-
terial on loan as a further aid ; Dr. A. S. Romer, Director, Dr. P.
J. Darlington, Jr., Curator of Insects, Dr. Herbert W. Levi,
Associate Curator of Arachnology, and Miss Nelda E. Wright,
Editor of Publications, all of the staff of the Museum of Com-
parative Zoology at Harvard College.

The problem of evolution and interspecific relationships in
the genus Micrathena are exceedingly interesting, often complex,
baffling, and difficult to trace. Females tend to exhibit a high

392 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

degree of abdominal spination involving many different patterns,
and often show strongly contrasting and variable color patterns.
The males, on the other hand, as adults usually lack definite
spines, exhibit much less gaudy coloration, are greatly reduced
in size, and possess a set of complex variations in the minute
structure of their secondary sexual organs and related parts.
The high degree of sexual dimorphism has made it extremely
difficult to match adult males with the proper females. More-
over, adult males are relatively infrequent in collections. When
young the sexes often resemble each other rather closely and
this fact can sometimes be used as an aid in matching the sexes
accurately.

F. Pickard-Cambridge (1904) included a total of thirty-one
species in his treatment of the genus Micrathena. One of these
(A. fericulum 0. P. -Cambridge) has been transferred to the
genus lldebaha Keyserling. Of the remaining thirty species,
five had both sexes fully recognized, seven were known only
from males, and the remaining nineteen were known only from
females.

So far as I have been able to determine up to the present time,
a total of forty species of the genus Micrathena have been more
or less definitely reported from Central America. Eight of these
were males unassociated with females; six species included both
sexes ; twenty-six species were known only from females. As will
be shown in the appropriate places in this paper, all four species
described as new by Chamberlin and I vie (1986) have been
synonymized with already known species. It now seems quite
certain that M. secies (Getaz) is really M. sexspinosa (Halm).
It is now generally known that M. comigera (0. P-Cam-
bridge) is the male of M. sexspinosa (Halm) and that M. longi-
cauda (0. P. -Cambridge) is the male of M. horrida (Tac-
zanowski). It has now been possible to complete the identifica-
tion of the male of M. schreibersi (Perty). M. clypeata (Walck.),
until recently considered to be distinctly a South American
species, is now known to be well established in Panama. In addi-
tion to the changes noted above, I have been obliged to recognize
and describe nine new species, thus bringing the revised list to
forty-three as now reported from Central America. Anions
these are twenty different kinds of males, nine of which are still
unassociated with the proper females. The complete list as the

CHICKERING : MICRATHENA IN CENTRAL AMERICA 393

species are understood at the present time may be given as
follows :

M. bimucronata (0. P.-Cambridge), 1899

M. brevipes (O. P.-Cambridge), 1890

M. catenulata F. P.-Cambridge, 1904

M. clypeata (Walck.), 1805

M. crassa (Keyserling), 1863

M. disjuncta sp. nov.

M. donaldi sp. nov.

M. duodecimspinosa (O. P.-Cambridge), 1890

M. fidelis (Banks), 1909

M. flaveola (C. L. Koch), 1839

M. funebris (Banks), 1898

M. furcula (O. P.-Cambridge), 1890

M. gladiola (Walck.), 1841

M. gracilis (Walck.), 1805

M. granulata F. P.-Cambridge, 1904

M. horrida (Taczanowski), 1873

M. inaequalis F. P.-Cambridge, 1904

M. insolita sp. nov.

M. macfarlanei sp. nov.

M. macilenta sp. nov.

M. militaris (Fabricius), 1775

M. mitrata (Hentz), 1850

M. mirifica sp. nov.

M. modica sp. nov.

M. molesta sp. nov.

M. parallela (O. P.-Cambridge), 1890

M. patruelis (C. L. Koch), 1839

M. petersi (Taczanowski), 1872

M. quadriserrata F. P.-Cambridge, 1904

M. saccata (C. L. Koch), 1836

M. sagittata (Walck.), 1841

M. schreibersi (Perty), 1833

M. serrata F. P.-Cambridge, 1904

M. sexspinosa (Hahn), 1822

M. spinulata F. P.-Cambridge, 1904

M. striata F. P.-Cambridge, 1904

M. subflava sp. nov.

M. subspinosa F. P.-Cambridge, 1904

M. trapa (Getaz), 1891

M. triserrata F. P.-Cambridge, 1904

M. uncata F. P.-Cambridge, 1904

M. vitiosa (O. P.-Cambridge), 1890

M. zilehi Kraus, 1955

394 BULLETIN: MUSE! M OF COMPARATIVE ZOOLOGY

Holotypes of all new species described in this paper will be
deposited in the Museum of Comparative Zoology at Harvard
College. 31. donaldi sp. nov. is named for my son who accom-
panied me on two very pleasant and productive collecting trips
to Panama. M. macfarlanei sp. nov. is named for Mr. D. Mac-
farlane, Commonwealth Institute of Entomology, with whom
I was very pleasantly associated during my period of work in
the British Museum (Natural History) in the summer of
li!58.

Genus MlCRATHElS A Sundevall, 1833

A comprehensive definition of the genus Micrathena Sunde-
vall is very difficult to draw up because of the great differences
which exist between males and females and even among the two
sexes themselves. For the present I am simply adopting what 1
consider to be the general understanding among araneologists at
the present time. I cannot do better than to accept the definition
essentially as given by Xteimoser (1917) in his treatment of the
whole genus as he understood it at the time of the publication
of his paper.

No attempt is made to include all citations which have ap-
peared in the literature on this genus. Only those which are
considered as particularly pertinent are given here. Those who
desire more extensive bibliographies are referred to Fioewer
(1912) and Bonnet (1957).

Because of unusual difficulties, no satisfactory key to the
females in this genus has been worked out. This is in part due
to the fact that puzzling variations in spination and some other
features occur quite frequently. The accompanying key to the
males should be an aid in identification of the different mem-
bers of this sex.

Key to the Species of Micrathena in Central America

Males

1. With a ventral hook on first coxa and a corresponding ridge and groove
on second femur {clypeata, disjuncta, duodetimspinosa, furcida,
macilenta, mitrata, parallela, patruelis) 12

la. Without any ventral hook on first coxa and without a corresponding
ridge and groove on second femur (brevipes, donaldi, gracilis,
granidata, horrida, miriflca, modioa, sagittata, sclireibersi, sexspmosa,
u neat a, vitiosa) 9

CHICKERING : MICRATHENA IN CENTRAL AMERICA 395

2. Carapace with three pairs of clearly defined dorsolateral foveae ....

clypeata, p. 400

2a. Carapace without three pairs of clearly defined dorsolateral foveae
(disjuncta, duodecimspinosa, furcula, macilenta, mitrata, parallela,
patruelis) 3

3. Abdomen at least twice as long as wide (duodecimspinosa, furcula, ma-

cilenta, mitrata, parallela) 4

3a. Abdomen less than twice as long as wide (disjuncta, patruelis) 7

4. With palpal tibia considerably wider than long (macilenta, mitrata,

parallela ) 5

4a. With palpal tibia not notably wider than long (duodecimspinosa, fur-
cula) 8

5. Abdomen with clearly evident spinules at posterior end (macilenta,

parallela) 6

5a. Abdomen with no clearly defined spinules at posterior end

mitrata, p. 435

6. Basal palpal tarsal hook nearly as broad at tip as long (Fig. 137) . .

parallela, p. 443

6a. Basal palpal tarsal hook not nearly as broad at tip as long (Fig. 104)
macilenta, p. 432

7. Abdomen nearly rectangular in outline (Fig. 143) . . . patmelis, p. 445
7a. Abdomen more rounded laterally ; not so nearly rectangular in outline

(Fig. 26) disjuncta, p. 403

8. Abdomen with definite tubercles at posterior end, indicating suppressed

spines duodecimspinosa, p. 408

8a. Abdomen without definite tubercles at posterior end, indicating sup-
pressed spines furcula, p. 416

9. Abdomen slender, at least twice as long as broad (brevipes, gracilis,

granulata, horrida, modica, sagittata, schreibersi, sexspinosa, uncata)

9a. Abdomen not so slender, about twice as long as broad or less than
twice as long as broad (donaldi, mirifica, vitiosa) 18

10. Abdomen with a caudal-like extension (Figs. 75, 86; gracilis, horrida)

10a. Abdomen without any caudal-like extension (brevipes, granulata, mo-
dica, sagittata, schreibersi, sexspinosa, uncata) 12

11. Base of bulb of palpal tarsus with a rounded knob fitting into a cor-

responding concavity in the tarsal hook (Fig. 76) . gracilis, p. 421

11a. Base of bulb of palpal tarsus without such a knob and corresponding

concavity as in gracilis horrida, p. 424

12. Palpal tarsal hook extended into a long, distally curved, sickle-like

structure (Fig. 180) sexspinosa, p. 456

12a. Palpal tarsal hook not extended into a distinctly sickle-like structure
(brevipes, granulata, modica, sagittata, schreibersi, uncata) 13

13. Abdomen constricted in the middle and rounded posteriorly (Fig. 161)

schreibersi, p. 452

396 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

13a. Abdomen not constricted in the middle nor smoothly rounded posteriorly
(brevipes, granulata, modica, sagittata, uncata) 14

14. Palpal tibia extended distally into a pair of sharp spines (Fig. 156)

sagittata, p. 449

14a. Palpal tibia not extended distally into a pair of sharp spines {brevipes,
granulata, modica, uncata) 15

15. Palpal tibia extended into a series of three bluntly rounded outgrowths

(Pig. 7) brevipes, p. 398

15a. Palpal tibia not extended into a series of three bluntly rounded out-
growths (granulata, modica, uncata) 16

16. Palpal tarsal hook a strongly chitinized and granulated plate (Pigs.

81,82) granulata, p. 423

16a. Palpal tarsal hook without a strongly chitinized and granulated plate;
tarsal hook tending toward the type more frequently found in the
genus (modica, uncata) 17

17. First femur with a group of modified ventral spines near middle of the

segment (Fig. 203) uncata, p. 466

17a. First femur without the group of modified spines near middle of seg-
ment modica, p. 439

18. Abdomen with four pairs of definite but small spines (Fig. 31)

donaldi, p. 405

18a. Abdomen without any definite spines (mirifica, vitiosa) 19

19. Palpal tarsal hook extended into a quadrate, concave, finely granulose

plate; with remains of three pairs of spines in the form of tubercles

on abdomen vitiosa, p. 466

19a. Palpal tarsal hook not extended into a quadrate, concave, finely granu-
lose plate; abdomen without visible remains of tubercles on abdomen
mirifica, p. 437

Micrathena bimucronata (0. P. -Cambridge) , 1899

(Figures 1-5)

Acrosoma bimucronatum 0. P. -Cambridge, 1899

M. bimucronata F. P.-Cambridge, 1904

M. bimucronata Petrunkevitch, 1911

M. bimucronata Reimoser, 1917

.1/. bimucronata Chickering, 1931

M. bimucronata Mello-Leitao, 1932

M. bimucronata Roewer, 1942

M. bimucronata Bonnet, 1957

Several specimens of this species in the British Museum (Natu-
ral History) from Guatemala have been examined and found to
agree well with specimens from Costa Rica and Honduras now
in the collections of the Museum of Comparative Zoology at Har-
vard College. The male is still unknown. One of the specimens

CHICKERING: MICRAT1IENA IN CENTRAL AMERICA

397

in the collection of the Museum of Comparative Zoology has been
selected as the hypotype from which the following facts have
been derived: Total length 5.85 mm.; the abdomen has a short
conical tubercle at each anterolateral corner (one specimen was
found to have these tubercles drawn out into short spines thus
illustrating the variability of the species) and a long stout spine
at each posterolateral angle (Figs. 1, 2) ; the carapace has a

External Anatomy of Micrathena

Figures 1-5, M. bimucronata

Figures 6-8, M. brevipes

Fig. 1. Abdomen of female, dorsal view.

Fig. 2. Posterior surface of abdomen to show appearance of spines.
Figs. 3-5. Epigynum; from below, posterior surface, and right lateral
view, respectively.

Fig. 6. Body of M. brevi'pes, dorsal view.
Fig. 7. Right palpal tarsus and tibia.
Fig. 8. Palpal tarsal hook, more enlarged.

well defined central fovea and two pairs of moderately clear
dorsolateral foveae ; the characteristics of the epigynum are
shown in Figures 3-5. The species is now known from Guatemala,
Costa Rica and Honduras.

398 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Micrathena brevipes (0. P. -Cambridge) , 1890

(Figures 6-8)

Acrosomal brevipes O. P.-Cambridge, 1890

A. brevipes Keyserling, 1892

M. brevipes F. P.-Cambridge, 1904

M. brevipes Petrunkevitch, 1911

M. brevipes Reimoser, 1917

M. brevipes Petrunkevitch, 1925

M. brevipes Chickering, 1931

if. brevipes Roewer, 1942

M. brevipes Bonnet, 1957

This species was reported from Honduras in 1931 (Chicker-
ing) but the specimens seen at that time are not now in the
collection and cannot, therefore, be re-examined for accuracy of
determination. My only opportunity to study the species care-
fully came during my period of work in the British Museum
(Natural History) in the summer of 1958. Two specimens are
now in the vial marked "type" but both abdomens are detached
and one may not belong to M. brevipes (O.P. -Cambridge).
Both cephalothoraces and attached parts are in good condition.
The abdomen (Fig. 6) is injured but appears to be similar to
that of the male of M . sexspinosa. The chief features of the pal-
pal tarsus and tibia are shown in Figures 7 and 8. Keyserling 's
description (1892) appears to be accurate and detailed. Total
length about 4.5 mm. The legs appear to be devoid of significant
spines. The first coxae lack the ventral hook and the second
femora lack the corresponding ridges and grooves. The female
remains unknown. The species appears to be widely distributed
in Central America and has been reported from Mexico, Guate-
mala, Honduras, and Panama.

Micrathena catenulata F. P.-Cambridge, 1904
(Figures 9-13)

M. catenulata Petrunkevitch, 1911
Al. catenulata Reimoser, 1917
M. catenulata Roewer, 1942
M. catenulata Bonnet, 1957

In the Nathan Banks collection in the Museum of Comparative
Zoology T have found several females from Mexico which agree
well with specimens in the Pickard-Cambridge collection in the

CIIICKERING : MICRATIIENA IN CENTRAL AMERICA

399

British Museum (Natural History). I also have two specimens
collected by myself in Honduras. Two males were found with
the females from Mexico but they are still considered to repre-
sent a separate species for reasons which will be given in some
detail later in this paper.

Female hypotype. Total length 6.695 mm. One difference be-
tween the hypotype and the holotype is in respect to abdominal
spines. F. P. -Cambridge stated that the "anterior marginal
angles" of the abdomen were prominent but did not bear spines.
Apparently this is a variable feature because spines occur in
these regions in some specimens but not in all. Short anterior
marginal spines are extended from the corners of the abdomen
in the hypotype. There are also two pairs of relatively large
robust spines in dorsal anterior and posterior positions as indi-
cated in Figure 9. It was also stated in the original descriptions

External Anatomy of Micrathena

Figures 9-13, M. eatenulata

Fig. 9. Dorsal view of body.

Fig. 10. Posterior spine and cusp ; lateral view from right side.
Figs. 11-13. Epigynum from below, in posterior view, and in profile from
right side, respectively.

that immature specimens have a short cusp below each posterior
dorsal spine. I have found that this also sometimes occurs in
mature females (Fig. 10). The head portion of the carapace is
only moderately raised. The sternum is not notably produced

400 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

posteriorly. The features of the epigynum are shown in Figures
11-13. Color in alcohol : the Mexican and Honduran specimens
are much lighter in color than the Guatemalan specimens avail-
able to F. P. -Cambridge. Apparently the black spots have
been reduced and the white parts greatly extended as indicated
in Figure 9.

Collection records. The specimens studied by F. P. -Cam-
bridge were from Guatemala. Other specimens available to me
in this study are from Escuintla, Chiapas, Mexico, with no date
of collection given, and Lancetilla, Honduras, July, 1929.

Micrathena clypeata ( Walckenaer ) , 1805
(Figures 14-20)

Epeira clypeata Walckenaer, 1805

M. clypeata Sundevall, 1833

M. clypeata, C. L. Koch, 1838

Plectana clypeata Walckenaer, 1841

M. clypeata Taczanowski, 1879

M. clypeata Simon, 1895

M. clypeata Petrunkevitch, 1911

11. clypeata Reimoser, 1917

M. planata Chambeiiin and Ivie, 1936. New synonymy

31. clypeata Chiekering, 1936

M. clypeata Roewer, 1942

M. planata Roewer, 1942. New synonymy.

M. clypeata Bonnet, 1957

M. planata Bonnet, 1957. New synonymy.

Until comparatively recently this species has been considered
strictly South American, but is now known to be well established
in Panama. Chamberlin and Ivie (1936) have given a detailed
description of the species under the name M. planata.

Female hypotypc. The extremely flattened form of the abdo-
men, the numerous smoothly chitinized dorsal abdominal spots,
together with the ten abdominal spines (Fig. 14) should make
identification certain. Figures 15-17 show the chief features of
the epigynum.

Male hypotype. Total length 3.9 mm. With body extremely
flattened (Fig. 18) ; with a complete but thin dorsal abdominal
shield. The first coxa has a distal, ventral, retrolateral hook and
the second femur has a corresponding groove and ridge near
its proximal end nearly dorsal in position but on the prolateral
surface. Palp : the tarsal hook has a characteristic form ; this

CHICKERING : MICRATIIENA IN CENTRAL AMERICA

401

and other features of the organ are shown in Figures 1!) and
20.

Collection records. All of my records of this species in
Panama have come from Barro Colorado Island, C. Z. The
female hypotype was taken in August, 1954 ; the male hypotype
in August, 1950. Two other males are in the collection : July,

External Anatomy of Microthena

Figures 14-20, M. clypeata

Fig. 14. Dorsal view of body of female.

Figs. 15-17. Epigynum from below, in posterior view, and in profile (more
enlarged), respectively.

Fig. 18. Dorsal view of body of male.

Fig. 19. Palpal tarsus of male.

Fig. 20. Palpal tarsal hook, retrolateral view (more enlarged).

402

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1924 (Banks) and July, 1936. Females have been taken from
June to August, 1934," 1936, 1950, and 1954. Chamberlin and
Ivie (1936) reported both sexes from my collection of 1928.

Micrathena cbassa (Keyserling) , 1863
(Figures 21-25)

Aerosoma crassum Keyserling, 1863

A. crassum Keyserling, 1892

M . crassa Simon, 1895

M. crassa Petrunkevitch, 1911

M. crassa Reimoser, 1917

M. crassa Petrunkevitch, 1925

M. crassa Roewer, 1942

If. crassa Bonnet, 1957

If. crassa Chickering, 1960

Petrunkevitch (1925) reported this species from Panama but
it has not appeared in my collections nor have I found it recorded

External Anatomy of Micrathena

Figures 21-25, If. crassa

Pig. 21. Abdomen, lateral view.
Fig. 22. Cephalothorax, lateral view.

Figs. 23-2.". Epigynum; from below, posterior view, and in profile, re-
spectively.

CHICKERING : MICRATHENA TN CENTRAL AMERICA 403

by any other worker. During the summer of 1958 I had an
opportunity to study the species in the Keyserling collection
in the British Museum (Natural History) and I have commented
on it elsewhere (1960). The most significant features of the
species may be given as follows: There is considerable variation
in the size of the females with the length varying from about
7.6 to 8.5 mm. ; head and thorax well separated by a deep trans-
verse groove ; the median thoracic fovea is clearly denned ; behind
the median fovea the thorax rises very sharply into a pronounced
gibbosity (Fig. 22) ; there are no clearly defined dorsolateral
foveae ; sternum only moderately convex ; abdomen with four
pairs of spines as shown in Figure 21 ; features of the epigynum
as shown in Figures 23-25. Collection records are restricted
to Colombia and Panama. The male is still unknown.

MiCRATHENA DISJUNCTA Sp. UOV.

(Figures 26-30)

Male holotype. Total length 3.705 mm. Carapace 1.755 mm.
long, 1.24 mm. wide opposite interval between second and third
coxae where it is widest ; .66 mm. tall shortly behind well defined
central fovea ; rises gently from just behind PME to this region
and then descends to posterior border.

Eyes. Eight in two rows as usual. Seen from above, posterior
row moderately recurved, anterior row strongly recurved. Seen
from in front, anterior row very slightly recurved, posterior row
gently procurved, all measured by centers. Ocular tubercles
moderately well developed. Central ocular quadrangle only
slightly wider behind than in front, slightly wider behind than
long. Ratio of eyes AME : ALE : PME* : PLE = 5:5:
6.5 : 5 (long diameters used when there are differences).
AME separated from one another by nearly two diameters,
from ALE by about three diameters. PME separated from
one another by nearly five-fourths of their diameter, from
PLE by nearly three times their diameter. Laterals only
separated by a broad line. Height of clypeus equal to a little
more than twice the diameter of AME.

Chelicerae, Maxillae, and Lip. All apparently normal to males
of the genus. Fragility of the specimen prevents close examina-
tion of such structures as teeth along the fang groove but details
appear to be unnoteworthy for the proper description of the
holotype.

404

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Sternum. Simple scutiform ; moderately convex ; terminates
bluntly between fourth coxae which are separated by about one-
half of their width.

Legs. 1423. Width of first patella at "knee" .16245 mm.,
tibial index of first leg 10. Width of fourth patella at "knee"
.11913 mm., tibial index of fourth leg 10.

External Anatomy of Micrathena

Figures 26-30, M. disjuncta

Fig. 2C. Dorsal view of body of male.
Fig. 27. Right first femur, prolateral view.
Fig. 28. Right second femur, prolateral view.
Fig. 29. Male palpal tarsus.
Fig. 30. Palpal tarsal hook, more enlarged.

1.540

.506

1.320

.440

.836

.264

CHICKERING : MICRATHENA IN CENTRAL AMERICA 405

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

.990 .924 .462 4.422

.72(i .748 .400 3.634

.440 .418 .330 2.288

4. 1.540 .396 .814 .770 .435 3.955

Palp .198 .0975 .1300 — .440 .866

With numerous robust spines on legs 1, 2, and 4 ; those on femora
appear to be most significant (Figs. 27-28). First coxa with a
ventral hook and second femur with the corresponding proximal,
prolateral ridge and groove.

Palp. Most essential features shown in Figures 29 and 30.
Both tibia and patella are very short and without special
modifications. Tarsal hook similar to that found in several other
species.

Abdomen General shape shown in Figure 26. Poorly devel-
oped tubercles at posterior end may represent suppressed spines
which show in immature males and in adult females.

Color in alcohol. Carapace yellowish white in a broad central
stripe with a broad dusky stripe on each side (Fig. 26). Legs
yellowish with variations. Mouth parts and sternum nearly
white. Abdomen: dorsum with a broad, central, broken, white
stripe and a semi-transparent cross in the middle ; nearly black
across posterior end and on each side a nearly black stripe (Fig.
26) ; just lateral to the black stripe are narrow, irregular,
brownish stripes.

Type locality. The holotype male is from Barro Colorado Is-
land, C. Z.. January, 1958. No paratypes have appeared in my
collection. The female is unknown.

MlCRATHENA DONALDI sp. UOV.

(Figures 31-34)

Male holotype. Total length 4.68 mm. Carapace 1.917 mm.
long; 1.125 mm. wide opposite second coxae where it is widest;
median eyes on a prominent tubercle and lateral eyes on two
confluent tubercles; only slightly raised behind median thoracic
fovea.

Eyes. Eight in two rows as usual; viewed from above, both
rows strongly recurved ; viewed from in front, both rows pro-
curved. Central ocular quadrangle wider behind than in front
in ratio of 25 : 17, wider behind than long in ratio of 25 : 22.

406 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Ratio of eyes AME : ALE : PME : PLE = 8 : 7 : 10 : 8. AME
separated from one another by five-eighths of their diameter,
from ALE by seven-fourths of their diameter. PME separated
from one another by nearly their diameter, from PLE by three-
halves of their diameter. LE separated from one another by
about the diameter of AME. Height of clypeus equal to nearly
twice the diameter of AME. Clypeus sharply slanted backward
because of strong development of median ocular tubercle.

Chelicerae. Moderately well developed ; parallel. Teeth along
fang groove not observed because of danger of injury to holo-
type.

Maxillae and Lip. As usual in males of the genus, without
noteworthy features.

Sternum. Moderately convex ; strongly and irregularly corru-
gated throughout ; sternal suture procurved ; with well developed
lateral sternal tubercles and low tubercles opposite coxae 1-3 and
with a blunt, low tubercle at posterior end which is not extended
between fourth coxae which are only slightly separated.

Legs. 4123. Width of first patella at "knee" .12996 mm.,
tibial index of first leg 10. Width of fourth patella at "knee"
.11913 mm., tibial index of fourth leg 7.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1. 1.170 .450 .845 .747 .520 3.732

2. 1.170 .410 .780 .650 .520 3.530

3. .780 .270 .520 .440 .396 2.406

4. 1.625 .390 1.200 1.105 .625 4.945
Palp .528 .176 .396 ■ .726 1.826

Leg spines very sparsely and poorly developed. Coxal ventral
hook and corresponding second femoral groove and ridge lack-
ing.

Palp. General characters like those of male of M. sagittata
but with tibia resembling that of M. brevipes (Figs. 32-34). The
tibia appears to be quite distinctive and the same can be said
for the tarsal hook.

Abdomen. General form shown in Figure 31. Abdominal
spines much more prominent than usual in mature males ; two
posterior pairs are relatively large and conspicuous; there is a
pair of short marginal spines somewhat behind the middle and
another pair of very small dorsal spines somewhat in front of
the middle : these suggest that the female should be found among

CIIK'KERING : MICRATIIENA IN CENTRAL A MKKK 'A

401

those with four pairs of more or less conspicuous abdominal
spines such as M. fidelis (Banks).

Fig.
Fig.
Fig.
ferent
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

External Anatomy of Micratliena

Figures 31-34, If. donaldi
Figures 35-40, M . duodecimspinosa

31. Dorsal view of body of male.

32. Left palpal patella, tibia, and tarsus, nearly prolateral view.

33. Distal end of palpal conductor and associated structures; dif-
view from that in Fig. 32.

34. Palpal basal tarsal hook.

35. Dorsal view of male allotype.

36. Eight second patella and tibia, prolateral view.

37. Left palpal tibia and tarsus.

38. Posterior surface of cymbium and tarsal hook from a paratype.

39. Palpal tibia and tarsal hook.

40. Another view of tarsal hook more enlarged ; from a paratype.

408 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Color in alcohol. Legs dark brown, almost black, above, but
yellowish below. Carapace and sternum a dark mahogany
brown. Abdomen : dorsum with three whitish spots as shown in
Figure 31 ; with remaining surface a mosaic of black, whitish,
brown, much streaked and dotted.

Type locality. Holotype male from Barro Colorado Island,
C. Z., August, 1936. Two paratype males from the same locality,
July, 1936. Mr. Banks collected one male in the same localitv,
July, 1924.

Micrathena duodecimspinosa (0. P. -Cambridge) , 1890

(Figures 35-45)

Acrosoma 12-spinosum O. P. -Cambridge, 1890

Acrosoma 12-spinosa Keyserling, 1892

M. 12-spinosa F. P.-Cambridge, 1904

A. 12-spinosa Banks, 1909

M. duodecimspinosa Petrunkevitch, 1911

M. duodecimspinosa Reimoser, 1917

M. duodecimspinosa Chiekering, 1936

M. duodecimspinosa Roewer, 1942

M. duodecimspinosa Kraus, 1955

M. duodecimspinosa Bonnet, 1957

Females of this species have been well described by Keyser-
ling (1892), F. P.-Cambridge (1904), and Reimoser (1917).
Abdominal spines are probably the best single clues to identifica-
tion.

Female hypotype. Total length 7.475 mm. The salient features
of body form are shown in Figures 41 and 42. In all of my
Panamanian specimens the posterior spines are bright red. Fea-
tures of the epigynum are shown in Figures 43-45.

Kraus (1955) has recently reported finding immature males
but, so far as I have been able to determine, the mature male
has never been described. For this reason, one of several males
from Boquete, Chiriqui, Panama, has been selected as the allo-
type and described as follows :

Male allotype. Total length 4.973 mm. Carapace 1.885 mm.
long, 1.43 mm. wide opposite interval between second and third
coxae where it is widest ; median fovea well developed ; nearly
level throughout from PME to posterior declivity (Fig. 35).

Eyes. Eight in two rows as usual ; viewed from above, both
rows strongly recurved ; viewed from in front, anterior row
slightly procurved and posterior row strongly procurved, all

CHICKERING : MICRATHENA IN CENTRAL AMERICA

409

measured by centers. Central ocular quadrangle wider behind
than in front in ratio of 10 : 9 ; wider behind than long in
about the same ratio. Ratio of eyes AME : ALE : PME : PLE
= 6 : 4.5 : 8 : 5. AME separated from one another by about
their diameter, from ALE by about 3.3 times their diameter.
PME separated from one another by about their diameter, from
PLE by 2.5 times their diameter. Laterals only slightly separ-
ated. Laterals on a rather marked tubercle. Height of clypeus
equal to about 1.5 times the diameter of AME.

External Anatomy of Micrathena
Figures 41-45, M. duodeeims'pinosa

Fig. 41. Dorsal view of body of female.

Fig. 42. Spines at posterolateral corner of abdomen; viewed from pos-
terior surface.

Figs. 43-45. Epigynum from below, posterior surface, and in profile
(right side), respectively.

Chelicerae. Moderately well developed ; essentially parallel ;
only slightly concave along outer border ; basal boss small but
clearly present ; apparently four teeth on promargin and three
on retromargin of well developed fang groove (observed on
paratype to avoid injury to allotype).

Maxillae and Lip. As usual in the genus; details unnote-
worthy.

Sternum. Only slightly convex; extended between all coxae
and with posterior coxae only slightly separated.

410 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Legs. 1423. Width of first patella at "knee" .18411 mm.,
tibial index of first leg 12. Width of fourth patella at "knee"
.16245 mm., tibial index of fourth leg 12.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1. 1.430 .435 1.105 1.072 .585 4.627

2. 1.300 .422 .845 .910 .520 3.997

3. .950 .325 .520 .550 .390 2.735

4. 1.820 .390 .910 .975 .500 4.595
Palp .352 .132 .120 .572 1.176

There is a rather poorly developed ventral distal hook on the
first coxa and a corresponding short prolateral proximal groove
and ridge on the second femur. The first two pairs of legs are
notably spiny, particularly the tibiae ; the ventral spines on the
second tibiae (Fig. 36) are short and robust. Trichobothria are
prominent on the tibiae.

Palp. Both patella and tibia are very short. The features of
the tarsus and tarsal hook are shown in Figures 37-40.

Abdomen. 3.120 mm. long; continued posterior to spinnerets
about four-ninths of total length ; remains of posterolateral
spines, prominent in females, appear as short tubercles at pos-
terior end. Other features as usual in the genus.

Color in alcohol. Legs brownish with variations; first two
darkest. Chelicerae and palps yellowish with palpal tarsus light
brown. Lip very dark brown like sternum. Maxillae dark gray-
ish. Carapace brownish with variations. Abdomen with a dark
irregular central series of spots and an irregular series of dorso-
lateral white spots making a pattern difficult to describe (Fig.
35) ; venter brownish dappled with yellowish white spots.

Collection records. The hypotype female is from Barro Color-
ado Island, C. Z., July, 1954. The allotype male is from Boquete,
Chiriqui, Panama, July, 1939 at which time several paratype
males were also taken. Females and immature males are in the
collection from Barro Colorado Island, C. Z., June, 1934; June-
July, 1936; June, 1939; July, 1954; January, 1958; France
Field, C. Z., August, 1939. Several females are in the collection
of the Museum of Comparative Zoology from Rio Jesus Maria,
Costa Rica, January (Biolley and Tristan), and also from San
Jose, Costa Rica (Valerio). The Cambridges had females from
Mexico, Guatemala, and Panama.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 411

MlCRATHENA FIDELIS (Banks), 1909

(Figures 46-49)

Acrosoma fidelis Banks, 1909
M. fidelis Petrunkevitch, 1911
Al. fidelis Eeimoser, 1917
il. fidelis Boewer, 1942
M. fidelis Bonnet, 1957

This species is, apparently, known only from the holotype fe-
male which was very briefly described by its author without
illustrations. These facts would seem to warrant a full descrip-
tion in accord with my usual formula. The description is taken
directly from the original specimen.

Female holotype. Total length 7.02 mm. Carapace 2.34 mm.
long; 2.015 mm. wide opposite second coxae where it is widest;
with the median fovea as usual in the genus ; cephalic portion
somewhat convex behind PME, then transversely depressed just
anterior to median fovea ; conspicuously swollen just posterior
to median fovea.

Eyes. Eight in two rows as usual ; lateral eyes on a slightly
raised tubercle ; viewed from above, both rows moderately re-
curved; viewed from in front, anterior row slightly recurved,
posterior row gently procurved ; central ocular quadrangle wider
behind than in front in ratio of 17 : 14, slightly wider behind
than long. Katio of eyes AME : ALE : PME : PLE = 4.5 :
5:7: 3.5. AME separated from one another by their diameter,
from ALE by four times their diameter. PME separated from
one another by about five-sevenths of their diameter, from PLE
by a little less than three times their diameter. Laterals almost
in contact. Height of clypeus equal to a little less than the diam-
eter of AME.

Chelicerae. Robust, vertical, nearly parallel, very convex in
front, especially in proximal half ; fang fairly robust and evenly
curved ; promargin of fang groove with four teeth, retromargin
with three.

Maxillae and Lip. As usual in the genus ; without noteworthy
features.

Sternum. Elongate scutiforni; quite convex throughout; not
continued between coxae ; with procurved sternal suture and
moderately well developed sternal tubercles at each anterolateral
corner. Posterior coxae separated by a little more than one-
seventh of their width.

412

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Legs. 1423. Width of first patella at "knee" .352 mm.,
tibial index of first leg 13. Width of fourth patella at "knee"
.264 mm., tibial index of fourth leg 12.

External Anatomy of Hierathena

Figures 46-49, M. fidelis
Figures 50-54, M. flaveola

Fig. 46. Dorsal view of body of female.

Fig. 47-49. Epigynum from below, in posterior view, and in profile from
right side, respectively.

Fig. 50. Dorsal view of body of female.

Fig. 51. Lateral view of spines at posterolateral corner of abdomen.

Figs. 52-54. Epigynum from below, in posterior view, and in profile from
right side, respectively.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 413

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

12.210

.975

1.800

1.495

.775

7.255

1.950

.855

1.365

1.280

.660

6.110

1.300

.455

.785

.775

.600

3.915

2.405

.715

1.430

1.495

.650

6.695

Legs with numerous spines and low tubercles from which hairs
extend. Numerous trichobothria observed on tibiae and meta-
tarsi.

Abdomen. General form and eight spines shown in Figure 46.
Otherwise as usual in the genus.

Epigynum. In general quite similar to others in the genus
but a profile view seems to show distinctive features (Figs.
47-49).

Type locality. The only specimen known to me is the holotype
described by Banks (1909) and collected at Tablazo, Costa Rica
(Tristan), September. The male remains unknown.

MlCRATHENA FLAVEOLA (C. L. Koch), 1839

(Figures 50-54)

Acrosoma flaveolum C. L. Koch, 1839
Plectana flaveola Walckenaer, 1841
M. flaveola Petrunkeviteli, 1911
11. flaveola Reimoser, 1917
M. flaveola Roewer, 1942
M. flaveola Bonnet, 1957

This species has caused me much concern and I still regard its
status in Central America as somewhat questionable. The species
has been reported from Mexico, Brazil, Dutch Guiana, and
Venezuela. I am convinced that it has often been confused with
M. gladiola (Walckenaer). Some of the specimens assigned to
this species and to M. gladiola may well belong to other species,
perhaps undescribed. I seem to have sound reasons for including
the species among the Central American forms treated in this
paper but I have had only one specimen for careful study and
that was not in prime condition. The following facts are taken
from this specimen :

Female hypotype. Total length to posterior border of abdo-
men between posterior spines 8.5 mm. (considerably larger than
described by previous workers). The general form of the body

414 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

is shown in Figures 50 and 51. The carapace is somewhat gib-
bous just behind the central fovea ; the sternum is finely rugulose
and moderately convex but is not raised into a distinct cone as
in M. gladiola (Walckenaer). The legs are badly fragmented
and unmeasurable in this specimen. The spines appear as shown
in Figures 50 and 51 and agree well with descriptions given by
other authors who have had numerous specimens with which to
work. The features of the epigynum are given in Figures 52-54.
Males are still unknown. The hypotype female is from San Jose,
Costa Rica, with no date of collection given.

Micrathena funebris (Banks), 1898
(Figures 55-59)

Acrosoma funebre Banks, 1898
M. funebris Petrunkeviteh, 1911
M. funebris Reimoser, 1917
M. funebris Chamberlin, 1924
M. fimebris Boewer, 1942
Al. funebris Bonnet, 1957

Only the female of this species is known but there are now
numerous specimens of this sex in the Museum of Comparative
Zoology representing parts of the Banks and Chamberlin collec-
tions. Two of the specimens in the Banks collection are labelled
"cotypes." In view of the brevity of the original description
and the lack of accompanying figures, one of the "cotypes" has
been selected and described in accord with my usual procedure.

Female hypotype. Total length 6.825 mm. Carapace 2.275 mm.
long, 1.592 mm. wide opposite second coxae where it is widest;
more slender than in many species (Fig. 55). Without additional
noteworthy features.

Eyes. Eight in two rows as usual; median eyes and laterals
on low tubercles ; viewed from above, both rows recurved ; viewed
from in front, anterior row gently recurved and posterior row
procurved, both measured by centers ; central ocular quadrangle
wider behind than in front in ratio of 4 : 3, wider behind than
long in ratio of 16 : 13. Ratio of eyes AME : ALE : PME :
PLE = 7.5 : 5.5 : 10 : 6. AME separated from one another by
eleven-sevenths of their diameter, from ALE by a little more
than four diameters. PME separated from one another by
about 1.5 times their diameter, from PLE by three diameters.

CHICKERING : MICRATHENA IN CENTRAL AMERICA

415

Laterals separated from one another by nearly the diameter of
ALE. Height of clypeus nearly equal to twice the diameter
of AME.

Chelicerae. Apparently as usual in the genus. Impossible to
view the fang groove without serious injury to specimen.

Maxillae. Without noteworthy features.

Lip. Wider than long in ratio of 10 : 7 ; deeply grooved in
basal fourth.

External Anatomy of Micrathena

Figures 55-59, M. funebris
Figure 60, M. furcula

Fig. 55. Body of female, dorsal view.
Fig. 56. Abdominal spines at posterolateral angle.

Figs, fw-59. Epigynum from below, from posterior view, and in profile
from right side, respectively.

Fig. 60. Body of female in dorsal view.

Sternum. Elongate scutiforni; longer than wide at anterior
end in ratio of 23 : 18 ; sternal suture proeurved ; with low
tubercles at anterolateral corners ; with a prominent tubercle at
posterior end ; not extended between fourth coxae which are
separated by about one-third of their width.

416 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Legs. 4123. Width of first patella at "knee" .255 mm., tibial
index of first leg 13. Width of fourth patella at "knee" .242
mm., tibial index of fourth leg 12.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1.755

.650

1.250

1.300

.650

5.605

1.625

.650

.975

1.235

.640

5.125

1.170

.325

.715

.835

.520

3.565

2.405

.585

1.430

1.235

.520

6.175

Apparently the leg spines were present in the usual manner but
are now mostly lacking.

Abdomen. The general form of this part of the body is best
shown in Figures 55 and 56. There are no anterior spines on
the abdomen but six are present as shown. The color pattern
appears to be characteristic and shows only minor variations
among the numerous specimens from islands in the Gulf of
California. Other features as usual in the genus.

Epigynum. Strongly tubercular; somewhat distinctive (Figs.
57-59).

Color in alcohol. In general, very dark brown or black with
a rather distinctive pattern of white spots on the dorsum ; essen-
tially as described by the author of the species and with little
change after long preservation.

Type locality. The original specimens came from Calmalli
Mines, Sierra San Nicolas, and Mazatlan, Mexico (Tristan).
The hypotype is from Mazatlan. There are many specimens from
islands in the Gulf of California in the Chamberlin collection.

Micrathena furcula (O. P. -Cambridge) ,1890

(Figures 60-66)

Acrosoma furcula O. P.-Cambridge, 1890

A. furcula Keyserling, 1892

M. furcula Simon, 1895

M. furcula F. P.-Cambridge, 1904

M. furcula Petrunkevitch, 1911

.1/. furcula Eeinioser, 1917

M. furcula Banks, 1929

M . furcula Eoewer, 1942

M. furcula Bonnet, 1957

CHICKERING : MICRATHENA IN CENTRAL AMERICA

417

Only the female has been known up to the present time. Now
however, with the finding of immature males with a copious sup-
ply of hair as well as other revealing features the male can be
positively identified. Banks was the first to report the female
from Panama.

Female hypotype. Total length 11.375 mm. Carapace quite
typical of the genus except that it is largely covered by a
copious growth of long yellowish silky hair. The abdomen is
also well supplied with hair but this is of a darker color and
largely confined to the lateral margins and especially to the
posterior furcula. The color as noted among the specimens avail-
able for study is quite variable but follows rather closely the

External Anatomy of Micrathena

Figures 61-66, If. furcula

Figs. 61-63. Epigynum from below, in posterior view, and in profile from
tight side, respectively.

Fig. 64. Body of male in dorsal view.
Fig. 65. Palpal patella, tibia, and tarsus.
Fig. 66. Palpal tarsal basal hook.

description given by the author of the species. The form of
the abdomen (Fig. 60) should make identification certain. The
characteristics of the epigynum are shown in Figures 61-63.

418 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Male allotype. Total length 4.94 mm. Carapace 2.145 mm.
long, 1.820 mm. wide opposite interval between second and third
coxae where it is widest ; without any dorsal swellings ; markedly
narrowed opposite first coxae (Fig. 64).

Eyes. Eight in two rows as usual ; ocular tubercles prominent,
especially the laterals; viewed from above, both rows recurved;
viewed from in front, anterior row gently recurved, posterior
row procurved ; central ocular quadrangle wider behind than in
front in ratio of 9 : 8, about as long as wide behind. Ratio
of eyes AME : ALE : PME : PLE = 5.5 : 4 : 6.5 : 3.5. AME
separated from one another by slightly more than their diameter,
from ALE by three diameters. PME separated from one another
by a little less than their diameter, from PLE by nearly three
diameters. Laterals separated from one another by nearly the
radius of AME. Height of clypeus equal to about 2.5 times the
diameter of AME ; clypeus very receding.

Chelicerae. With low basal tubercle in front ; unable to observe
fang groove and teeth without serious injury to specimen.

Maxillae. Without noteworthy features.

Lip. Wider at base than long in ratio of 4 : 3 ; transversely
and shallowly grooved ; with sternal suture procurved.

Sternum. Elongate scutiform ; widest between second coxae;
longer than wide in ratio of about 3:2; considerably and irregu-
larly corrugated ; narrowly extended between fourth coxae which
are only slightly separated.

Legs. 4123. ' Width of first patella at "knee" .17328 mm.,
tibial index of first leg 12. Width of fourth patella at "knee"
.15162 mm., tibial index of fourth leg 11.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1.592

.400

1.085

1.105

.575

4.757

1.495

.390

.845

1.000

.520

4.250

.975

.260

.520

.515

.390

2.660

1.950

.400

.975

.520

4.820

Palp

.400

.110

.608

1.228

There is a moderately well developed ventral distal retrolateral
hook on the first coxa and the corresponding short prolateral
groove and ridge on the second femur. The first two legs are
notably spiny, especially on the ventral and prolateral surfaces
of the tibiae but the detailed placement seems not to be essential
to the adequate description of the allotype. The ventral spines

CHICKERING : MICRATHENA IN CENTRAL AMERICA 419

on the two anterior tibiae are short and robust. Trichobothria
are conspicuous on the tibiae.

Palp. Both patella and tibia are short with the latter relatively
broad ; the shape of the basal tarsal hook is shown in Figure
66 ; other tarsal features are shown in Figure 65.

Abdomen. With a moderately well developed dorsal scutum;
2.9 mm. long; 1.46 mm. wide in middle; only moderately flat-
tened; other features fairly typical of males of the genus (Fig.
64).

Color in alcohol. Carapace a rich medium brown with fine
black dots ; legs generally yellowish brown but the first two pairs
of femora are darker brown like the carapace. Sternum brown,
mottled with black. Abdomen: dorsum with a color pattern
essentially as represented in Figure 64. The darkly colored areas
are a deep brown, nearly black; the lighter areas are yellowish
beneath the scutum but white around the margin ; laterally there
is a series of seven white elongated spots ; the venter is a complex
of irregular blackish and brown spots and streaks. The color
pattern in mature males is probably quite variable. As already
pointed out, one important clue to proper identification of the
male is the copious supply of hair in immature specimens.

Type locality. The holotype female came from Guatemala.
The hypotype female is from Barro Colorado Island, C. Z., July,
1936. The allotype male is from Barro Colorado Island, C. Z.,
July, 1934. Two immature males are in my collection from
the same locality, June, 1939. Several mature females have been
taken on Barro Colorado Island, C. Z., on the following dates:
July, 1924 (Banks). July-August, 1936.

Micrathena gladiola (Walckenaer). 1841
(Figures 67-71)

Acrosoma aculeata C. L. Koeh, 1836 (preoccupied)

Plcctana gladiola Walckenaer, 1841

A. flaveolum Keyserling, 1892

M. gladiola F. P.-Cambridge, 1904

M. gladiola Petrunkevitch, 1911

M. gladiola Reimoser, 1917

M. fl areola Petrunkevitch, 1925

.1/. gladiola Roewer, 1942

M. gladiola Bonnet, 1957

In collections made available to me I have found much con-
fusion regarding the identification of this species. I have found

420

bulletin: museum of comparative zoology

it assigned to other species and other species assigned to it. I
am convinced that the group of species to which this belongs
should be carefully studied, especially after more careful collect-
ing is done within its area of distribution. In view of the uncer-
tainties surrounding the species, I am relying quite largely upon
the descriptions and identifications of both F. P. -Cambridge
and Keyserling. The most significant features of the species,
taken from a specimen collected at Bugaba, Panama, appear to
be the following : The carapace is uniformly convex, not raised
behind the median fovea ; lateral eyes are on a low tubercle ; fang
groove with four teeth on promargin and three on retromargin;
sternum very convex, raised into a distinct cone ; epigynum with
features as shown in Figures 69-71 taken from a specimen

External Anatomy of Miorathena

Figures 67-71, M . gladiola

Fig. 67. Abdomen of female in dorsal view.

Fig. 68. Abdominal spines at posterolateral angle from left side.
Figs. 69-71. Epigynum from below, in posterior view, and in profile from
right side, respectively.

studied by F. P. -Cambridge in the British Museum (Natural
History) ; femora finely granulose and with minute setigerous
granules in front and beneath ; abdominal spines as shown in
Figures 67 and 68. The species is quite well known from a few
places in Central America, northern South America, and some
of the West Indies nearest to South America. The male is still
unknown.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 421

IMtcrathena gracilis ( Walekenaer) , 1805
(Figures 72-77)

Epeira gracilis Walekenaer, 1805

Pie ei ana gracilis Walekenaer, 1841

Acrosomal matronale C. L. Koch, 1845

E. rugosa Hentz, 1850

A. rugosum Emerton, 1884

M icrathena gracilis Simon, 1895

M. matronalis Simon, 1895

.1/. gracilis F. P. -Cambridge, 1904

A. gracilis Banks, 1909

M. gracilis Petrimkevitch, 1911

M. gracilis Reimoser, 1917

M. gracilis Chickering, 1931

M. nigrior Chamberlin and Ivie, 1936. New synonymy.

M. gracilis Eoewer, 1942

H. gracilis Kraus, 1955

M. gracilis Bonnet, 1957

This species is widely distributed from the northern United
States through Central America to Brazil. It has been figured
and described many times but males remain scarce in collections
and there is some evidence of confusion of these with males of
other species. Only two males have been found thus far in the
collections from Central America. The only record from Panama
is that of M . nigrior Chamberlin and Ivie, 1936. This is reported
to have come from my collection of 1928 but it seems odd that it
has not appeared in any of my numerous collections taken since
that date.

Female hypotype. Total length 13 mm., including the prom-
inent bases of the chelicerae. Cephalic part of the carapace cpiite
prominent : both lateral ocular tubercles and median ocular
tubercles moderately prominent ; central ocular quadrangle only
slightly wider behind than in front, longer than wide behind in
ratio of 8 : 7. Sternum with a pointed tubercle at posterior end.
Femora densely granulose with setigerous tubercles. Fang
groove with four teeth along the promargin and three along the
retromargin. Abdomen very tall; with five pairs of spines (Fig.
72) which should aid greatly in identification. Epigynum more
distinctive than usual in the genus (Figs. 73-74).

Male hypotype. Total length 6.11 mm. Abdomen extremely
long and slender; strongly chitinized; with a pair of segmental
divisions at posterior end, the second of which bears a pair of

422

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

stout terminal spines (Fig. 75). The legs appear to lack the
modified spines so frequently present in the males of the genus.
The first coxa lacks the ventral hook and the second femur also
lacks the ridge and groove so frequently present. The chief
palpal features are shown in Figures 76-77.

Collection records. The female hypotype is from Patulul, Gua-
temala, January, 1912 (W. M. Wheeler). The male hypotype is
from Orizaba, Mexico, with no date given. Other female speci-
mens studied by me are from: Granada, Nicaragua (C. F.
Baker); Polvon, Nicaragua; Tampico, Mexico, 1913 (H. L.
Locke); Vera Cruz, Mexico; San Jose, Tamaulipica, Mexico,
July, 1930 (Bartlett and Dice) ; Uricuajo, Costa Rica (Biolley
and Tristan). One record only from Panama: M. nigrior Cham-
berlin and Ivie.

External Anatomy of Micrathena

Figures 72-77, M. gracilis

Fig. 72. Abdomen of female, dorsal view.

Figs. 73-74. Epigynum in posterior and profile views, respectively.

Fig. 75. Dorsal view of body of male.

Fig. 76. Left palpal tarsus.

Fig. 77. Palpal tarsal basal hook, more enlarged.

(.MUCKERING: MICRATIIENA IN CENTRAL AMERICA

423

Micrathena granulata F. P. -Cambridge, 1904

(Figures 78-82)

^[. granulata Petrunkevitch, 1911
M. granulata Eeimoser, 1917
M. granulata Boewer, 1942
.1/. granulata Bonnet, 1957

Male hypotype. Total length 4.16 mm. General form as shown
in Figure 78. Legs with ordinary and unnoteworthy spination
except for the first femora which, apparently, have a group of
clasping spines near the distal end (Fig. 79). The first coxa

External Anatomy of Micrathena

Figures 78-82, M. granulata

Fig. 78. Dorsal view of body of male.
Fig. 79. Distal end of left first femur; ventral view.

Figs. 80-82. Three different views of left palpal tibia and tarsus showing
denticulated basal tarsal hook.

lacks a ventral hook and the second femur lacks the correspond-
ing ridge and groove. Palp: the tarsal hook is very distinctive,
being broadly extended, concave, apically recurved, and set with
many minute denticles (Figs. 80-82). Color in alcohol: Legs,

424 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

cephalothorax, and mouth parts all light reddish brown with
variations ; abdomen dorsally yellowish with irregular whitish
patches (Fig. 78).

For a time it was thought that M. granulata was the missing
male of M. catenulata. Recently a male belonging to this species
was found with females clearly belonging to M. funebris (Banks)
collected on Coronados Island, Gulf of California. At present
it would seem that 31. granulata F. P. -Cambridge could be the
missing male for either M. catenulata F. P. -Cambridge or M.
funebris (Banks). On the other hand, perhaps this species be-
longs with some other of the numerous species known only from
females. This condition again emphasizes the need for careful
collecting and close observation over the whole area occupied
by this genus.

Collection records. F. P. -Cambridge had the species only from
Teapa, Mexico. The male hypotype is from Chiapas, Escuintla,
Mexico with no date of collection given. I have a second speci-
men taken with the hypotype and another from Coronados Is-
land, Gulf of California, May 18, 1921. These three are all in
the collection of the Museum of Comparative Zoology at Harvard
College.

^ov

Micrathena horrida ( Taczanowski) , 1873
(Figures 83-88)

Acrosoma horrida Taczanowski, 1873

A. mammiUata Butler, 1873

A. longicauda Keyserling, 1892. New synonymy.

M. mammiUata F. P. -Cambridge, 1904

A. longicauda O. P. -Cambridge, 1890. New synonymy.

M . longicauda F. P. -Cambridge, 1904. New synonymy.

A. mammiUata Banks, 1909

M. horrida Petrunkevitch, 1910

M. horrida Petrunkevitch, 1911

H. mammiUata Petrunkevitch, 1911

M. longicauda Petrunkevitch, 1911. New synonymy.

M. horrida Reimoser, 1917

M. longicauda Reimoser, 1917. New synonymy.

M. horrida Petrunkevitch, 1925

M. longicauda Petrunkevitch, 1925. New synonymy.

M. longicauda Banks, 1929. New synonymy.

A. mammiUata Banks, 1929

31. mammiUata Bryant, 1940

M. horrida Roewer, 1942

CHICKERING : MICHATHENA IN CENTRAL AMERICA

425

M. longicauda Koewer, 1942. New synonymy.

M. horrida Bonnet, 1957

M. longicauda Bonnet, 1957. New synonymy.

Until recently it has seemed unsafe to American araneologists
to synonymize M. mammillata (Butler) with M. horrida (Tacz-
anowski). It now seems to me that the synonymy as indicated is

External Anatomy of Micrathena

Figures 83-88, M. horrida

Fig. 83. Dorsal view of body of female.

Figs. 84-85. Epigynum in posterior view, and in profile, right side, re-
spectively.

Fig. 86. Dorsal view of body of male.

Fig. 87. Left palpal tibia and tarsus of male.

Fig. 88. Male palpal tibia and basal tarsal hook to show form of latter.

426 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

fully justified. F. P. -Cambridge suggested that 31. longicauda
(0. P.-Cambridge) might be the male of M. mammillata (But-
ler). Banks (1929) was quite convinced that this was so but he
continued to list them separately. Miss Bryant (]940) recog-
nized the male of M. mammillata but did not synonymize it with
M. longicauda. The species appears to be closely related to M.
gracilis (Walck.) ; this conclusion is based upon a comparison
of both sexes of both species.

Female hypotype. Total length 8.9 mm., including the prom-
inent bases of the chelicerae. Very strongly chitinized. Carapace
with raised cephalic part ; only slightly raised behind median
fovea. Sternum convex but without a posterior tubercle as seen
in M. gracilis. The femora are excessively provided with seti-
gerous tubercles. Abdomen : with seven pairs of marginal spines
the first of which extends from the anterior margin over the
posterior lateral corners of the carapace (Fig. 83) ; there are
also numerous lateral tubercles beneath the marginal spines (five
on each side in the hypotype) subject to considerable variation
among the many specimens available ; the epigynum is strongly
tubercular (Figs. 84-85) ; there is also a prominent, median,
chitinized tubercle just posterior to the cone surrounding the
spinnerets.

Male hypotype. Total length 4.85 mm. Very long and slender ;
strongly chitinized (Fig. 86). Abdomen with a trilobate pos-
terior termination ; the trilobate termination bears four small but
definite spines. Legs without especially modified spines; femora
one and two with numerous small setigerous tubercles ; patellae
one and two quite dilate on retromargin ; coxa one without any
ventral hook and femur two without the corresponding ridge
and groove. Features of the palp with tarsal hook shown in Fig-
ures £7-88. The shape of the abdomen and the palpal features
furnish the best means of identification of the male of the species.

Collection records. Male and female hypotypes are from Barro
Colorado Island, C. Z., July and August, 1950, respectively. The
species is widely distributed from the southern United States
through Central America to several countries in northern South
America and the West Indies. It occurs in my collections from
many localities in Panama and is abundant on Barro Colorado
Island.

CHICKERING : MICRATHENA IN CENTRAL AMERICA

427

Micrathena inaequalis F. P. -Cambridge, 1904

(Figures 89-93)

Acrosoma inaequalis Banks, 1909
M. inaequalis Petrunkevitch, 1911
M. inaequalis Beimoser, 1917
M. inaequalis Chickering, 1936
M. inaequalis Boewer, 1942
M. inaequalis Bonnet, 1957

F. P. -Cambridge (1904) had this species from Costa Rica and
Guatemala. Banks (1909) had it from Costa Rica. I reported
it from Panama in 1936 but at that time the specimens in my
possession were all immature and may have been wrongly iden-
tified. Now I can definitely record the species from the high-
lands of western Panama.

External Anatomy of Micrathena

Figures 89-93, M. inaequalis

Fig. 89. Dorsal view of body of female.
Fig. 90. Posterolateral abdominal spines; posterior view.
Figs. 91-93. Epigynum from below, in posterior view, and in profile from
right side, respectively.

Hypotype female. Total length 7.8 mm. Carapace raised both
before and behind the median fovea. Sternum moderately con-
vex ; with a series of low marginal tubercles. Abdomen : there

428 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

is a pair of sharply pointed spines reaching far over the cara-
pace ; there is also a pair of short marginal spines somewhat
behind the middle ; the abdomen extends far laterally at the
posterior border and there terminates in a pair of spines on
each side (Figs. 89-90). These eight spines seem to be typical
of the species but small lateral marginal spines may also occur
on some specimens. The degree to which the large posterior lat-
eral spines are separated is also, apparently, at least somewhat
variable in the general population. Characteristics of the epigy-
num are shown in Figures 91-93.

Collection records. In addition to the earlier records of
the species from Costa Rica and Guatemala I can now add
the following from Panama : El Valle, July, 1936 ; Boquete.
Chiriqui, July, 1939; El Volcan, Chiriqui, August, 1950. The
hypotype fe2iiale is from Boquete, July, 1939. The male is un-
known.

MlCRATIIEXA INSOLITA Sp. 110V.

(Figures 94-98)

Apparently the holotype had recently completed its last moult
and was not yet well chitinized. It is, however, in good condi-
tion for describing.

Female holotype. Total length 4.55 mm. Carapace largely
covered by the anterior marginal abdominal spines and project-
ing anterior end of the abdomen itself ; about 2.08 mm. long and
about 1.625 mm. wide opposite interval between second and
third coxae where it is widest; median thoracic fovea obscure;
regularly and gently arched dorsally from PME to posterior
declivity with no marked gibbosity as in many species in the
genus.

Eyes. Eight in two rows as usual ; viewed from above, both
rows recurved ; viewed from in front, both rows procurved, pos-
terior row strongly so ; central ocular quadrangle wider behind
than in front in ratio of about 3 : 2, wider behind than long in
ratio of about 18 : 13. Ratio of eyes AME : ALE : PME : PLE
= 7 : 7 : 8 : 7 (laterals somewhat oval). AME separated from
one another by a little more than 1.5 times their diameter, from
ALE by nearly six times their diameter. PME separated from
one another by slightly more than 2.5 times their diameter, from
PLE by about 5 times their diameter. Laterals separated from
one another by about the radius of ALE. Height of clypeus equal
to 1.5 times the diameter of AME.

(MUCKERING : MICRATIIENA IN CENTRAL AMERICA

429

Chcliccrae. Parallel, moderately robust ; as usual in the genus.
Unable to see fang- groove because of fragility of the holotype.

Maxillae and Lip. Apparently as usual in the genus and with-
out noteworthy features.

External Anatomy of Micrathena

Figures 9-4-98, M. insolita

Fig. 94. Dorsal view of body of female.

Fig. 95. Posterior end of abdomen ; seen from posterior surface.
Figs. 9(i-98. Epigynum from below, from a somewhat posterior view, and
in profile from right side, respectively.

Sternum. Quite convex; without tubercles; posterior coxae
separated by about half their width.

Legs. 4123. Width of first patella at "knee" .2274 mm., tibial
index of first leg 12. Width of fourth patella at "knee" .2373
mm

tibial index of fourth leg 13.

Femora Patellae Tibiae Metatarsi
(All measurements in millimeters)

Tarsi

Totals

1.690

.715

1.170

1.040

.617

5.232

1.660

.715

1.040

.975

.520

4.910

1.170

.650

.530

.747

.520

3.617

2.145

.650

1.170

1.202

.617

5.784

430 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Legs with few spines; details of spination seem to be unnote-
worthy.

Abdomen. Measured with anterior and posterior spines, as
long as entire body. Three pairs of spines as indicated in Fig-
ures 94-95. Perhaps the most striking feature is the extreme
extension of the anterior pair of spines.

Epigynum. Features of this organ essentially as shown in Fig-
ures 96-98. This organ lies so close to the chitinized cone sur-
rounding the spinnerets that it is impossible to view it from
the posterior surface in the usual manner.

Color in alcohol. Legs generally yellowish ; first two pairs with
a dorsal brown stripe on femora, patellae, tibiae, and metatarsi
with the tarsi entirely brown ; second two pairs of legs similarly
colored except the brown stripe is changed to a large spot of
irregular shape. Carapace yellowish in anterior half with brown
dots and a narrow brown median stripe; posterior half with
brown dots closely crowded together. Abdomen: dusky yellowish
dorsally with black stripes and spots as suggested in Figure 94 ;
posterior and lateral surfaces yellowish with irregular black
stripes.

Type locality. Female holotype from Porto Bello, Panama,
August, 1936. There are no paratypes and the male is unknown.

MlCRATHENA MACFARLANEI Sp. UOV.

(Figures 99-102)

Female holotype. Total length from AME to middle of pos-
terior margin of abdomen 11.05 mm. Carapace about 4 mm. long
(considerably overlapped by abdomen), 3.185 mm. wide opposite
posterior border of second coxae where it is widest ; quite strongly
gibbous just posterior to well defined median fovea ; with paired
dorsolateral foveae very faintly indicated ; gently raised just
posterior to PMB.

Eyes. Eight in two rows as usual ; viewed from above, both
rows moderately recurved; viewed from in front, anterior row
slightly recurved, posterior row slightly procurved, all measured
by centers. Central ocular quadrangle wider behind than in
front in ratio of about 19 : 17 ; wider behind than long in ratio
of about 19 : 16. Ratio of eyes AME : ALE : PME : PLE =
6.5 : 4.5 : 7 : 4.5. AME separated from one another by ten-thir-
teenths of their diameter, from ALE by about 4 times their
diameter. PME separated from one another by slightly more
than their diameter, from PLE by four times their diameter.

CHICKERING : MICKATIIENA IN CENTRAL AMERICA

4:.U

Laterals separated from one another by slightly less than one-
fourth of the diameter of one of them. Height of elypeus equal
to slightly more than the diameter of AME.

Chelicerae. Robust; gibbons in front near base. Promargin of
fang groove with four teeth, relromargin with three robust teeth.

Maxillae and Lip. Typical of females of the genus; details
regarded as unnoteworthy.

Sternum. Sternal suture procurved; anterolateral tubercles
moderately well developed, others hardly tubercular, more
rounded ridges ; continuous posteriorly with a sclerite extending
between fourth coxae which are separated by two-fifths of their
width.

cT%E\

External Anatomy of Micrathena

Figures 99-102, M. macfarlanei

Fig. 99. Dorsal view of body of female.

Figs. 100-102. Epigynum from below, in posterior view, and in profile
from right side, respectively.

Legs. 4123. Width of first patella at "knee" .520 mm., tibial
index of first leg 10. Width of fourth patella at "knee" .520 mm..

tibial index of fourth leg 12.

432 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

5.005

1.365

3.575

1.105

14.625

4.615

1.300

3.055

3.120

.975

13.065

2.730

.925

1.625

1.480

.845

7.605

6.500

1.240

3.315

4.355

1.235

16.645

Setigerous tubercles numerous on legs; true spines few; first
femora with a series of six or seven very short prolateral spines.

Abdomen. Plump and full ; somewhat convex dorsally ; general
form and five pairs of spines as shown in Figure 99.

Epigynum. Essentially as shown in Figures 100-102.

Color in alcohol. Legs with varying shades of reddish brown.
Carapace with a narrow, median, dusky brown stripe from be-
tween PME to near median fovea; with a broad, broken, dusky
brown, dorsolateral stripe on eacli side leaving the margins yel-
lowish. Sternum dark brown. Lip dark brown, nearly black,
with a yellowish tip. Maxillae dark brown with anterior and
median margins yellowish. Chelicerae : fang and tips of basal
segments dark brown, elsewhere yellowish. Abdomen: dorsal
region almost white with black ' ' punctures ' ' ; second, fourth, and
fifth spines reddish at bases with more or less black striping;
lateral sides and venter a variegated black and yellowish.

Type locality. Female holotype from Barro Colorado Island,
C. Z., August, 1954; one mature paratype female and several
immature females from the same localitv, July, 1936 and June,
1939.

MlCRATHENA MACILENTA Sp. 110 V.

(Figures 103-109)

Note : The two specimens treated under the name M. macilenta
sp. nov. were formerly believed to belong to the species named
M. parallela (O. P.-Cambridge) but a careful comparison with
the type of the latter species in the British Museum (Natural
History) has shown this to be an error and so they must be
regarded as representing a species new to science. One is selected,
therefore, as the holotype and described in accord with my
usual procedure.

Male holotype. Total length 3.95 mm. Carapace 1.76 mm. long;
somewhat overlapped by abdomen ; 1.30 mm. wide slightly behind
second coxae where it is widest; .66 mm. tall just behind well
developed median fovea where it is tallest.

CHICKERING : MICRATHENA IN CENTRAL AMERICA

433

Eyes. Eight in two rows as usual ; ocular tubercles only mod-
erately developed; viewed from above, both rows strongly re-
curved; viewed from in front, anterior row slightly procurved,
posterior row strongly procurved, all measured by centers. Cen-
tral ocular quadrangle wider behind than in front in ratio of
11 : 9, wider behind than long in ratio of 22 : 19. Ratio of
eyes AME : ALE : PME : PLE = 7:6:8: 5.5. AME sep-
arated from one another by slightly less than their diameter,

External Anatomy of Micratliena

Figures 103-109, M. macilenta

Fig. 103. Dorsal view of body of male.

Fig. 10-4. Left palpal tarsus.

Fig. 105. Left first femur; prolateral view.

Fig. 106. Distal end of left second femur and patella ; prolateral view.

Fig. 107. Dorsal view of left palpal tibia.

Figs. 108-100. Two views of left palpal cymbium and basal tarsal hook.

from ALE by nearly two and one-third times their diameter.
PME separated from one another by their diameter, from PLE
by two and one-half times their diameter. Laterals separated
from one another only by a well defined line. Height of clypeus
equal to about one and one-fifth times the diameter of AME.

434 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Chclicerae, Maxillae, and Lip. All apparently quite typical of
males of the genus. Teeth along fang groove not observed because
of fragility of the holotype.

Sternum. Only slightly convex; slightly rugulose; without
definite tubercles; continued as a narrow sclerite between coxae
laterally and as a fairly broad, thin sclerite between fourth
coxae which are separated by about one-third of their width.

Legs. 1423. Width of first patella at "knee" .17328 mm.,
tibial index of first leg 11. Width of fourth patella at "knee"
.15162 mm., tibial index of fourth leg 13.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1.520

.528

1.056

1.000

.484

4.588

1.300

.440

.814

.770

.440

3.764

.704

.290

.396

.418

.352

2.160

1.540

.352

.814

.980

.418

4.104

Palp

.308

.141

.572

1.162

The ventral, distal, retrolateral hook on first coxa and the cor-
responding ridge and groove on the proximal prolateral surface
of the second femur are moderately well developed. There are
numerous spines on legs; those with most significance appear to
be on the first and second femora essentially as shown in Figures
105-106.

Palp. General features as shown in Figures 104, 107-109. Re-
sembles that of M. parallela (O. P. -Cambridge) with which the
species was formerly confused.

Abdomen. Much flattened dorsoventrally ; the only evidence
of suppressed spines shows at posterior end where two pairs of
small tubercles occur ; other features as usual in males of the
genus (Fig. 103).

Color in alcohol. Legs and mouth parts variable shades of
yellowish ; not distinctive. Carapace yellowish in the center,
brownish elsewhere. Abdomen : dorsum nearly white throughout
center with irregular black marginal stripes as shown in Figure
103; venter dusty yellowish with gray patches in regions of
booklungs. Paratype follows the color pattern of the holotype
very closely.

Type locality. Male holotype from Barro Colorado Island,
C. Z., July, 1934; one paratype male from the same locality,
August, 1950. and one taken in December, 1957. The female is
unknown.

CHECKERING : MICRATHENA IN CENTRAL AMERICA 435

Micrathena militabis (Fabricius) , 1775

Aranea militaris Fabricius, 1775

A. militaris Olivier, 1789

Plectana militaris WaJckenaer, 1841

Acrosoma militaris Butler, 1873

A. mil Hare Banks, 1898

M. militaris Petrunkevitch, 1911

.1/. militaris Eeimoser, 1917

M. militaris Petrunkevitch, 1930

M . militaris Eoewer, 1942

M. militaris Bonnet, 1957

Banks (1898), with some uncertainty, reported this species
from Margarita Island, Lower California. The single specimen
now in the Museum of Comparative Zoology from Margarita
Island and labelled M. militaris (Fabr.) is a specimen of M.
sexspinosa (Halm). Another specimen from Western Mexico
with a similar label is a female of M. sagittata (Walck.). I am
assuming, therefore, that M. militaris (Fabr.) has not yet been
found in Central America.

Micrathena mitrata (Hentz), 1850

(Figures 110-116)

Acrosoma mitrata Hentz, 1850

A. mitrata Emerton, 1884

A. reduvianum McCook, 1893

A. mitrata Banks, 1898

A. mitrata Emerton, 1902

M. mitrata F. P. -Cambridge, 1904

M. reduviana Petrunkevitch, 1911

M. mitrata Eeimoser, 1917

H. mitrata Eoewer, 1942

M. mitrata Kaston, 1948

.1/. mitrata Levi, 1954

M. mitrata Bonnet, 1957

The female of this species is well known from many localities
in the United States. It has also been reported from Mexico and,
doubtfully, from Cuba (Franginillo). The male, apparently, has
not been carefully studied and appears to be scarce in collections.
In general, the specimens from the United States studied by me
agree well with those from Mexico in the British Museum (Nat-
ural History). The hypotypes have been selected from collec-
tions made in Canton, North Carolina, and kept in the Museum

436

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

of Comparative Zoology. There is no date of collection given
but the specimens have been in preservation a long time.

Female hypotype. Total length 4.55 mm. Probably the best
means of identification is the presence of two pairs of postero-
lateral abdominal spines (Fig. 110). Prominent spines are lack-
ing elsewhere but there are many small lateral spinules usually

115

External Anatomy of Hicrathena

Figures 110-116, M. mitrata

Fig. 110. Dorsal view of body of female.

Figs. 111-113. Epigynum from below, in posterior view, and in profile
from right side, respectively.

Fig. 114. Dorsal view of body of male.

Fig. 115. Left palpal tarsus of male.

Fig. 116. Palpal tibia and basal tarsal hook of male, more enlarged.

CHICKERING : MICRATIIENA IN CENTRAL AMERICA 437

not mentioned in descriptions. The lateral spinules appear to be
less evident in the specimens from Mexico than among those
from the United States which I have had an opportunity to
study. In some of the specimens in the British Museum (Natu-
ral History) from Mexico there appeared to be a tendency for the
anterolateral corners of the abdomen to be somewhat extended.
The epigynum is obscurely distinctive (Figs. 111-113).

Male hypotype. Total length 3.25 mm. Abdominal spines are
completely suppressed in the male (Pig. 111). Features of the
palp are difficult to determine and present in drawings because
of long preservation, minute size, and some deterioration, but
Figures 115-116 are believed to give the most important charac-
teristics. The coxal hook and corresponding femoral groove and
ridge are poorly developed.

MlCRATIIENA MIRIFICA sp. 110V.

(Figures 117-121)

Male holotype. Total length 4.03 mm. Carapace 1.69 mm.
long; 1.495 mm. wide opposite interval between second and third
coxae where it is widest ; considerably overlapped by anterior
end of abdomen ; median fovea only slightly behind middle ;
median region nearly level from PME to posterior declivity ;
with lateral margins regularlv rounded to cephalic region (Fig.
117).

Eyes. Eight in two rows as usual; viewed from above, both
rows recurved ; viewed from in front, anterior row nearly
straight, posterior row procurved, all measured by centers. Cen-
tral ocular quadrangle only slightly wider behind than in front,
slightly longer than wide behind. Ratio of eyes AME : ALE :
PME : PLE = 7:5:7:5 (laterals somewhat angular). AME
separated from one another by slightly less than their diameter,
from ALE by about 2.5 times their diameter. PME separated
from one another by slightly less than their diameter, from PLE
by about three times their diameter. Laterals separated from
one another by about one-third of their diameter. Height of
clypeus equal to about 1.5 times the diameter of AME. Clypeus
quite receding. Median ocular tubercle quite prominent ; lateral
ocular tubercles moderately prominent.

Chelicerae, Maxillae, and Lip. Quite as usual in males of the
genus and without noteworthy features.

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Sternum. Nearly flat ; without noteworthy features. Pedicel
with pointed tubercle projecting forward toward posterior end
of sternum. Posterior coxae separated by a little more than half
their width.

Legs. 1423. Width of first patella at "knee" .166 mm., tibial
index of first leg 10. Width of fourth patella at "knee" .154
mm., tibial index of fourth leg 14.

Femora Patellae Tibiae Metatarsi
(All measurements in millimeters)

Tarsi

Totals

1.852

.540

1.040

.525

4.997

1.430

.410

.715

.780

.500

3.835

.760

.300

.455

.400

.325

2.240

1.625

.390

.745

.845

.455

4.060

Palp

.325

.152

.162

.638

1.277

Spines: first femur without special clasping spines; first tibia
with long, robust, prolateral, and ventral spines as shown in
Figure 120; second tibia with short, robust, prolateral, and
ventral spines as shown in Figure 121. Without coxal hook
and femoral ridge and groove.

117

118

External Anatomy of Miorathena

Figures 117-121, M. mirifiea

Fig. 117. Dorsal view of body of male.

Fig. 118. Left palpal tibia and tarsus.

Fig. 119. Left palpal patella, tibia, and basal tarsal hook.

Figs. 120-121. Left first and second tibiae, respectively; ventral view.

CIIICKERING : MICRATIIENA IN CENTRAL AMERICA 439

Palp. The tibia is simple, without spines or special modifica-
tions. Other palpal features, including tarsal hook, shown in
Figures 118-119.

Abdomen. Wholly without spines; considerably flattened dor-
soventrally; with form as shown in Figure 117.

Color in alcohol. Legs yellowish brown above with small dusky
patches ; lighter below. Carapace light brown in center with
broad, darker brown lateral stripes. Abdomen nearly white dor-
sally with black markings as suggested in Figure 117. Sternum
yellowish with a large black spot in posterior half. Color pattern
of paratypes like that of holotype.

Type locality. Male holotype and one paratype from Barro
Colorado Island, C. Z., July, 1950; one additional paratype from
the same locality, July, 1939.

MlCRATIIENA MODICA sp. 110Y.

(Figures 122-126)

Male holotype. Total length 4.29 mm. Carapace 1.625 mm.
long; 1.43 mm. wide just behind second coxae w7here it is widest;
thoracic part regularly rounded laterally; .585 mm. tall just
behind the moderately well marked central fovea where it is
tallest ; considerably overlapped by anterior border of abdomen.

Eyes. Eight in two rows as usual ; lateral ocular tubercles mod-
erately well developed; central ocular tubercle hardly existent.
Viewed from above, both rows rather strongly recurved ; viewed
from in front, anterior row gently recurved, posterior row
clearly procurved, all measured by centers. Central ocular quad-
rangle only slightly wider behind than in front, almost exactly
as long as wide behind. Katio of eyes AME : ALE : PME :
PLE =8 : 6 : 8.5 : 6. AME separated from one another by
nearly their diameter, from ALE by about three-halves of their
diameter. PME separated from one another by about their
diameter, from PLE by about twice their diameter. Laterals
separated from one another by a broad line. Height of elypeus
equal to about nine-eighths of the diameter of AME.

CheHcerae, Maxillae, and Lip. Apparently quite typical of
males in the genus. Teeth along the fang groove not observed
because of fragility of the holotype.

Sternum. Only slightly convex; tubercles almost completely
suppressed ; posterior end extended between fourth coxae which
are separated by about two-thirds of their width.

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BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Legs. 1243. Width of the first patella at "knee" .19494 mm.,
tibial index of first leg 9. Width of fourth patella at "knee"
.15162 mm., tibial index of fourth leg 9.

Femora

Patellae

Tibiae

Metatarsi

Tarsi

Totals

(All

measurements in

millimeters)

2.470

.600

1.690

1.625

.650

7.035

2.085

.575

1.320

1.365

.520

5.865

1.170

.357

.585

.720

.390

3.222

1.975

.455

1.170

1.276

.540

5.416

Palp

.286

.132

.154

.528

1.100

The ventral distal retrolateral hook on first coxa and the corre-
sponding proximal prolateral groove and ridge on the second
femur are all lacking in this species. The legs are all quite spiny
but the modified spines appear to be most important on the
first femur (Fig. 123). Probably a considerable degree of
variation in the spination will be noted when a series of these
males is available for study.

122

External Anatomy of Micrathena
Figures 122-126, M. modica

Dorsal view of body of male.

Left first femur ; ventral view.
Fig. 124. Left palpal tibia and base of tarsus with tarsal hook.
Fig. 125. Left palpal tibia and tarsus.
Fig. 126. Another view of basal tarsal hook.

Fig
Fig

i oo

123.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 441

Palp. General features shown in Figures 124-126. The hasal
tarsal hook resembles those of certain other species but is also
somewhat distinctive. The tibia is conservative and without
special features.

Abdomen. General features as shown in Figure 122. Much
flattened dorsoventrally ; with no indication of suppressed spines.

Color in alcohol. Legs and mouth parts in varying shades of
brownish and yellowish colors; details regarded as unessential.
Carapace with a yellowish brown central stripe and a broad
dusky brown stripe on each side as indicated by stippling in
Figure 122. Abdomen : with many white subchitinous flecks on
dorsum together with dark markings also as indicated in Figure
122.

Type locality. Male holotype is from Summit, Canal Zone,
July, 1950. One paratype male from Barro Colorado Island,
C. Z., July, 1950 ; one immature male from El Volcan, Chiriqui,
August, 1950 showing short spines at posterior end of abdomen.
The female is unknown.

MlCRATHENA MOLESTA sp. nOV.

(Figures 127-132)

Note : The holotype specimen described below was found in the
Nathan Banks collection in the Museum of Comparative Zoology.
It was filed with two other specimens and labelled M . triserrata
F. P. -Cambridge. I have carefully compared the holotype with
those specimens assigned to M. triserrata F. P. -Cambridge in
the British Museum (Natural History) and have been compelled
to decide that they do not belong together. For this reason, but
somewhat hesitantly, I have been compelled to consider the species
new to science and, therefore, I am describing the holotype in
accord with my usual procedure.

Female holotype. Total length 6.825 mm. Carapace about 2.21
mm. long; 1.82 mm. wide opposite the interval between second
and third coxae where it is widest; 1.40 mm. tall at level of
marked gibbosity just behind the central fovea which is well
defined ; with three pairs of dorsolateral foveae also well defined
(Fig. 127).

Eyes. Eight in two rows as usual ; viewed from above, pos-
terior row moderately recurved, anterior row strongly so. Viewed
from in front, anterior row gently recurved, posterior row gently
procurved, all measured by centers. Central ocular quadrangle

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BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

wider behind than in front in ratio of about 6 : 5, wider behind
than long in ratio of 15 : 14. Ratio of eyes AME : ALE : PME :
PLE = 9.5 : 7.5 : 11 : 6. AME separated from one another by
a little less than their diameter, from ALE by a little less than
2.5 times their diameter. PME separated from one another by
a little more than their diameter, from PLE by about five-thirds
of their diameter. Laterals separated from one another by about
one-fourth the diameter of PLE (a deformity has removed the
PLE far away from the ALE on the right side). Height of
elypeus equal to about the radius of AME.

131

External Anatomy of Micrathena

Figures 127-132, M. molettta

Fig. 127. Dorsal view of body of female.
Fig. 128. Posterior end of abdomen from behind.
Fig. 129. Eight lateral view of posterolateral corner of abdomen.
Figs. 130-132. Epigynum from below, in posterior view, and in profile
from right side, respectively.

Chelicerae, Maxillae, and Lip. All apparently quite typical of
the genus and with details regarded as unnoteworthy.

Sternum. A simple scutiform; sternal suture nearly straight;
anterolateral tubercles moderately developed, others hardly indi-
cated; not continued between fourth coxae which are separated
by about one-third of their width.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 443

Legs. 4123. Width of first patella at "knee" .22743 mm.,
tibial index of first leg 11. Width of fourth patella at "knee"
.23826 mm., tibial index of fourth leg 13.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters )

2.080

.725

1.430

1.105

.630

5.970

1.950

.655

1.190

1.040

.585

5.420

1.300

.425

.780

.650

.390

3.545

2.470

.640

1.430

1.495

.630

6.665

With few spines on legs ; exact number and placement not re-
garded as essential for adequate description ; with many seti-
gerous tubercles.

Abdomen. General form as shown in Figures 127-129. The
holotype has eight pairs of spines as shown ; one of the paratypes
has the same number but the other has the three small lateral
spines on each side represented only by tubercles.

Epigymim. Essentially as shown in Figures 130-132. One
paratype agrees well with the holotype in this respect but the
other does not, perhaps because of immaturity.

Color in alcohol. Carapace a reddish brown with lateral sides
dusky as indicated by stippling in Figure 127. Sternum a lighter
reddish brown. Legs and mouth parts with varying shades of
reddish brown. Abdomen : dorsum almost entirely yellowish with
small white subchitinous flecks; the numerous "punctures" are
reddish brown.

Type locality. Holotype female together with two paratype
females from Tablazo, Costa Rica. Apparently collected by Prof.
Tristan during the early part of the present century and re-
ported as M. triserrata F. P. -Cambridge (Banks, 1909). The
male is unknown.

Micrathena parallela (O. P. -Cambridge) , 1890
(Figures 133-137)

Acrosoma parallelum O. P. -Cambridge, 1890

A. parallelum Keyserling, 1892

M. parallela F. P.-Cambridge, 1904

M. parallela Petrunkevitch, 1911

M. parallela Reimoser, 1917

M. parallela Petrunkevitch, 1925

M. parallela Roewer, 1942

M. parallela Bonnet, 1957

444

BULLETIN: MUSEUM OE COMPARATIVE ZOOLOGY

The Pickard-Cambridges (1890, 1904) reported this species
from Panama and Guatemala. Keyserling (1892) mentions hav-
ing numerous specimens from Guatemala. The species has not
yet appeared in my collections and my only opportunity to study
it came during my period of work in the British Museum (Natu-
ral History) in the summer of 1958. The type material now
consists of parts of four specimens from which the following
facts have been taken. Apparently there are no highly modified
femoral spines in this species such as frequently occur in males
of the genus ; some variation in respect to tibial spines has been
noted among the available specimens but the ventral spines on
the first and second tibiae are probably fairly typical (Figs. 134-
135) ; there is some discoloration from long preservation but in

133

External Anatomy of Micrathena

Figures 133-137, M. parallela

Fig. 133. Dorsal view of body of male.

Fig. 134. Ventral spines of first tibia.

Fig. 135. Ventral spines of second tibia.

Fig. 13G. Palpal tarsus.

Fig. 137. Base of palpal tarsus with another view of basal tarsal hook.

general the color is a dull reddish brown with carapace lighter
in head region and along central region ; abdomen is irregularly

CIIICKERING : MICRATHENA IN CENTRAL, AMERICA 445

yellowish through the center and much darker on lateral sides ;
the body is much flattened dorsoventrally with lateral sides
nearly parallel ; the first coxa has the ventral hook well developed
and the second femur is provided with the corresponding groove
and ridge ; Figures 136-137 show the most important features of
the palpal basal tarsal hook and other parts of the palpal tarsus.
The female is unknown.

MlCRATHENA PATRUELIS (C. L. Koch), 1839

(Figures 138-145)

Acrosoma patruele C. L. Koch, 1839

Plectana patruela Walekenaer, 1841

P. reduviana Walekenaer, 1841

M. reduviana Simon, 1895

M. patruelis F. P. -Cambridge, 1904

M. reduviana F. P. -Cambridge, 1904

A. patruele Banks, 1909

M. patruelis Petrunkevitch, 1911

M. patruelis B-eimoser, 1917

M. patruelis dickering, 1931

.]/. retracta Chambeiiin and Ivie, 1936. New synonymy

M. patruelis Boewer, 1942

M. patruelis Bonnet, 1957

It is quite apparent that much confusion concerning this
species has existed in the past and I fear it is not yet entirely re-
moved. I am quite certain that M. retracta Chamberlin and Ivie
is the same as the species regarded as M. patruelis (C. L. Koch),
but I am not entirely satisfied that the latter, as now understood,
is one species alone. Kraus (1955) has recently identified M.
saceata (C. L. Koch), 1836 among his collections from El Salva-
dor. It seems to me that this raises the whole question of rela-
tionships between these two species and it is my opinion that
all available collections of both species should be carefully
studied in an endeavor to clarify the whole matter. At present I
cannot do better than to consider all specimens which I have
had an opportunity to study as belonging to M. patruelis (C. L.
Koch) as I have indicated.

Female hypotype. Total length 6.305 mm. Carapace only
slightly raised behind median thoracic fovea ; lateral margins
with numerous setigerous tubercles. Sternum moderately convex;
lateral tubercles extending toward intervals between second and
third coxae, third and fourth, and both fourth coxae quite

446

BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

marked. Abdomen with a pair of very short anterolateral tuber-
cles where spines so often occur in many species; with a pair of
posterolateral spines on each side with the upper one robust and

External Anatomy of Micratlwna

Figures 138-145, M. patruelis

Fig. 138. Dorsal view of body of female.
Fig. 139. Abdominal spines at posterolateral corner.

Figs. 140-142. Epigynum from below, in posterior view, and profile from
right side, respectively.

Fig. 143. Dorsal view of body of male.

Fig. 144. Palpal tibia and tarsus of male.

Fig. 145. Palpal tibia and basal tarsal hook in different view.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 447

projecting dorsally and the lower one much smaller and directed
posteriorly (Figs. 138-139) ; lateral margins and more dorsal of
the abdominal spines with minute denticles. Epigynum as shown
in Figures 140-142. Color in alcohol : Carapace and legs with
varying shades of brown ; sternum dark brown ; abdomen nearly
white above because of subchitinous granules but with dark
brown or black margins; venter and lateral sides dark brown
with many lighter stripes. Considerable variation has been noted
among available specimens in respect to color pattern. General
appearance shown in Figure 138.

Male hypotype. Total length 3.77 mm. First and second
femora with long spines above but only a terminal pair below ;
with many setigerous tubercles. First and second tibiae with
numerous modified spines. Only one prolateral spine on first and
second patellae as compared to two on each as mentioned by
F. P. -Cambridge. The usual coxal hook and femoral groove
and ridge are present. Palpal features are shown in Figures
144-145.^

Collection records. The species has been reported from Mexico,
through Central America and into northern South America. 1
have it for study from several parts of Panama, Honduras,
Nicaragua, Costa Rica, and Mexico. The male and female hypo-
types are both from Barro Colorado Island, C. Z., August, 1936
and July, 1954, respectively.

Micrathena petersi (Taczanowski) , 1872

Aorosoma petersi Taczanowski, 1872

A. petersi Keyserling, 1898

M. petersi F. P. -Cambridge, 1904

M. petersi Petrunkevitch, 1911

M. petersi Reimoser, 1917

M. petersi Banks, 1929

.1/. petersi Roewer, 1942

M. petersi Bonnet, 1957

Taczanowski described this species from Guiana. Keyserling
recorded it from Guatemala and described it from an immature
female. 1 have many immature specimens of M. sexspinosa
(Hahn) which closely resemble Keyserling 's figures and descrip-
tion. The specimens identified by Banks (1929) are all immature
and, according to my present view, all belong to M. sexspinosa
(Hahn). For these reasons I am not regarding M. petersi (Tac-
zanowski) as a Central American species.

448

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Micrathena quadriserrata F. P. -Cambridge, 1904
(Figures 146-149)

M. quadriserrata Petrunkevitch, 1911
M. quadriserrata Reimoser, 1917
M. quadriserrata Banks, 1929
M. quadriserrata Roewer, 1942
M. quadriserrata Bonnet, 1957

F. P. -Cambridge (1904) had the original specimens from
Guatemala; Banks (1929) recorded it from Panama; I have col-
lected it in Panama on several occasions. It has been reported
from South America but there seems to be some uncertainty
about the exactness of the identifications from this part of the
world. The male remains unknown as an adult.

Female hypotype. Total length 6.37 mm. The carapace is very-
gibbous behind the median fovea. The sternum is slightly convex.
Abdomen : there is a pair of long anterior spines extending far
over the carapace ; two pairs of small lateral marginal spines ;
the abdomen is bifurcated posteriorly and each bifurcation is
subdivided into four fairly robust spines (Fig. 146). The most
important epigynal features are shown in Figures 147-149. The

External Anatomy of Micrathena

Figures 146-149, M. quadriserrata

Fig. 146. Dorsal view of body of female.

Figs. 147-149. Epigynum from below, in posterior view, and in profile
from right side, respectively.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 449

most common color pattern is suggested in Figure 146; the un-
stippled areas are white and the stippled areas are black or gray ;
numerous specimens show that the color pattern is highly vari-
able ; in some the whole dorsum is black. In a specimen in the
British Museum (Natural History), collected in Joinville, Brazil,
there were a total of nine pairs of spines because of the multipli-
cation of small lateral spines to four on each side. The number
and placement of spines in Panamian specimens seem to be quite
consistent.

I have several immature males in my collection which I am,
for the present, regarding as the missing males in this species
but none is sufficiently mature to warrant description.

Collection records. The species has been recorded from both
Guatemala and Panama in Central America and from Brazil and
Venezuela in South America. I have taken the species repeatedly
on Barro Colorado Island, C. Z., at France Field, and at Fort
Sheridan, C. Z.

MlCRATHENA SACCATA (C. L. Koch), 1836

Acrosoma saccatum C. L. Koch, 1836

Plectana saccata Walckenaer, 1841

M. saccata Simon, 1895

M. saccata Petrunkevitch, 1911

M. saccata Beimoser, 1917

M. saccata Boewer, 1942

M. saccata Kraus, 1955

M. saccata Bonnet, 1957

Until recently this species has been considered exclusively
South American in distribution. Kraus (1955) reported it from
several localities in El Salvador. Of course, caution is indicated
here but it seems probable that the species has been confused
with M. patruelis (C. L. Koch) which it very closely resembles
and which is probably very common in El Salvador. For these
reasons I am not regarding the species as definitely established
in Central America.

Micrathena sagittata ( Walckenaer ) , 1841
(Figures 150-156)

Plectana sagittata Walckenaer, 1841
Epeira spinea Hentz, 1850
Acrosoma bovinum Thorell, 1859
A. bovinum Thorell, 1868

450 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

A. spineum Emerton, 1884

A. spineum McCook, 1893

A. spineum Emerton, 1902

M. sagittata F. P.-Cambridge, 190-4

M. sagittata Petrunkeviteh, 1911

.1/. sagittata Reimoser, 1917

M. sagittata Petrunkeviteh, 1930

M. sagittata Roewer, 1942

M. sagittata Kaston, 1948

M. sagittata Levi, 1954

M. sagittata Bonnet, 1957

This species has been figured and at least partially described
many times but the male is not easily separated from that of
several other species. M. sagittata is widely distributed through-
out the United States, Central America, West Indies, and the
northern parts of South America.

Female hypotype. Total length 8.45 mm. The cephalic portion
of the carapace is prominently raised ; considerably gibbous
behind the median fovea. The sternum is quite convex ; with
three pairs of lateral tubercles ; the posterior end is strongly
tubercular between the bases of the fouith coxae. Abdomen: with
a pair of long anterior spines arising from dorsolateral positions
(not anterior border) and extending far forward but not closely
contiguous to the carapace ; a pair of short lateral, somewhat re-
curved spines arise a little behind the middle ; a pair of long-
robust posterolateral spines ; each of the latter has a small cusp
at its base (often lacking in certain specimens) (Fig. 150). The
epigynum is more distinctive than usual ; the free part of the
scape appears very different than in most species when viewed
in profile (Figs. 151-153). Color in alcohol: dorsum of abdomen
largely yellowish with the long spines reddish at bases and black
at tips; lateral sides of abdomen with broken stripes of yellow
and black ; the cephalothorax is reddish brown with a conspicu-
ous whitish yellow margin.

Male hypotype. Total length 4.5 mm. The shape of the body
seems to vary somewhat among numerous specimens but the
shape of the hypotype is shown in Figure 154 and is regarded
as fairly typical. The color is dark brown dorsally with obscure
whitish spots as indicated by the unstippled areas. Palp : the
tibia is provided with two robust spines, a feature not usually
shown in descriptions or published figures ; the tarsal hook is
rather distinctive (Figs. 155-156). There is no ventral coxal
hook on the first coxa and no proximal prolateral groove or

CHICKERING : MICRATHENA IN CENTRAL, AMERICA

451

ridge on the second femur (eight males from North Carolina
and Colombia examined). Some specimens show the abdominal
spines, so conspicuous in females, as plainly suppressed spines in
the form of tubercles thus changing the form of the abdomen
considerably. This was especially true of the South American
forms.

1 5 O 151

External Anatomy of Micrathena

Figures 150-156, M. sagittata

Fig. 150. Dorsal view of body of female.

Figs. 151-153. Epigynum from below, in posterior view, and in profile from
right side, respectively.

Fig. 154. Dorsal view of body of male.
Fig. 155. Left palpal tibia and tarsus.
Fig. 156. Left palpal tibia and basal tarsal hook in different, view.

Collection records. F. P. -Cambridge had the species from
Mexico and Guatemala. Specimens from Mexico and Costa Rica
are in the collection in the Museum of Comparative Zoology at

Petrunkevitch (1930) has recorded it from

Harvard College

452 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Puerto Rico. I found numerous specimens from northern South
America in the British Museum (Natural History). It is well
known from many parts of the United States. It has not, how-
ever, appeared in my Panamanian collections. The male hypo-
type is from Balsam Gap, North Carolina, August, 1930; the
female hypotype is from Vera Cruz, Medias Aquas, Mexico, with
no date of collection given.

Micrathena SCHREIBERSI (Perty), 1833
(Figures 157-168)

Acrosoma schrcibersi Perty, 1833

A. spinosum C. L. Koch, 1836

Plectana macracantha Walckenaer, 1841

M. schrcibersi Simon, 1895

M. schrcibersi F. P.-Cambridge, 1904

.1/. schrcibersi Petrunkevitch, 1911

M. schrcibersi Beimoser, 1917

A. schreibersi Banks, 1929

M. coleophora Chamberlin and Ivic, 1936. New synonymy.

M. schrcibersi Boewer, 1942

M. schrcibersi Bonnet, 1957

This is a well known and strikingly beautiful species charac-
teristic of South America but now known to be one of the most
common members of the genus in Panama. Banks, apparently,
had the male properly identified but he did not report it in his
paper (1929). M. coleophora Chamberlin and Ivie is, quite
plainly, the male of this species.

Female hypotype. Total length, including the posterior abdom-
inal spines and the somewhat protruding bases of the chelicerae,
14.3 mm. The size appears to be quite variable as determined
from a study of a large number of individuals. There are ten
spines on the abdomen (Fig. 157). The head portion is strongly
raised and the carapace is moderately swollen behind the median
fovea. The sternum is only moderately convex with the anterior
border swollen into a low transverse ridge. The epigynum is
quite distinctive (Figs. 158-160). The color in alcohol is fairly
typical of the species in general ; the legs are a rich dark brown ;
the carapace is a similar brown with a yellowish-white margin ;
the abdomen is yellowish-white in the middle of the dorsum with
nearly black margins ; the anterior spines are white with the
second pair nearly black ; the fourth pair is bright red with
black tips ; the posterior pair is nearly black. The color pattern,

CIIICKERING : MICRATIIENA IN CENTRAL AMERICA

453

External Anatomy of Micrathena

Figures 157-160, M. schreibersi

Fig1. 157. Dorsal view of body of female.

Figs. 158-160. Epigynum from below, in posterior view, and in profile
(right side and somewhat more enlarged), respectively.

however, is highly variable as noted in a large series. One speci-
men from Colombia, for example, had bright yellow legs and a
very dark body.

Male hypotype. Total length 5.395 mm. The shape and gen-
eral appearance (Fig. 161) should serve to identify the males
of this species without much difficulty. The color is reddish
brown with variations. The abdomen is rounded behind and con-
stricted near the middle. Ventral spines on the first and second
tibiae are shown in Figures 167-168. Palp : the tibia appears to be
rather distinctive in shape ; the basal tarsal hook and other im-
portant features of the palpal tarsus are shown in Figures 162-
166. There is no ventral hook on the first coxa and no corre-
sponding groove and ridge on the second femur.

Collection records. The species has been reported from numer-
ous localities in South America, Mexico, and Panama. It is
probably widely distributed through Central America. It ap-
pears to be one of the two most common species of the genus on

454

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Barro Colorado Island, C. Z., but is much less common elsewhere
in Panama where I have collected. Both sexes are in my collec-
tion from Barro Colorado Island taken from June to August,
1934, 1936, 1939, 1950, 1954 and also in January 1958. I also
have it from C. Z. Forest Reserve, 1939: France Field, C. Z.,
August, 1939 ; Peluca Hydrographic Station, Boqueron River,
Panama, July, 1950.

External Anatomy of Micrathena
Figures 161-168, M. scJireibersi

Fig. 161. Dorsal view of body of male.

Figs. 162-163. Left palpal tarsus and basal tarsal hook; two different
views.

Fig. 164. Basal palpal tarsal hook ; seen from distal end of tarsus.

Figs. 165-166. Two different views of left palpal patella and tibia.

Figs. 167-168. Eight first and second tibiae, respectively, showing spines;
seen in ventral view.

CHICKERING : MICRATHENA IN CENTRAL AMERICA

455

Micrathena serrata F. P. -Cambridge, 1904
(Figures 169-173)

M. serrata Petrunkevitch, 1911
M. serrata Beimoser, 1917
M. serrata Checkering, 1936
.If. serrata Boewer, 1942
M. serrata Bonnet, 1957

This species seems to be very uncommon in collections. I had
my first opportunity to study the species carefully during my
period of work in the British Museum (Natural History) in the
summer of 1958. The species was reported from Barro Colorado
Island in my collection of 1934 (1936) but the specimens are not
now in the collection and, hence, the correctness of the identifica-
tion cannot now be determined.

Female hypotype. The following facts are taken from a cotype
in the British Museum (Natural History) : Total length from

External Anatomy of Micrathena

Figures 169-173, H. serrata

Fig. 169. Dorsal view of body of female.

Fig. 170. Spines at posterolateral corner of abdomen; right side.
Figs. 171-173. Epigynum from below, and in posterior view, and in profile
from right side, respectively.

456 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

chelicerae to posterior end of abdomen (exclusive of posterior
spines) 7.47 mm.; length to tip of posterior spines 8.07 mm.
Carapace 2.28 mm. long, 2.08 mm. wide at level of interval be-
tween second and third coxae where it is widest. Height of
clypeus slightly less than diameter of AME. Carapace consider-
ably raised behind conspicuously rounded central fovea ; with
three pairs of dorsolateral foveae (Fig. 169). Viewed from in
front, anterior row of eyes slightly recurved, posterior row
slightly procurved. Chelicerae with four teeth along promargin
of fang groove and three along retromargin. Sternum only
slightly convex; with posterior end slightly tubercular. Abdo-
men: extends forward over the carapace a moderate extent;
general form and seven pairs of spines as shown in Figures 169-
170. Features of the epigynum shown in Figures 171-173. The
species would seem to be closely related to M. duodecimspinosa
(0. P. -Cambridge). The male is unknown.

Collection records. The original specimens were all from the
highlands of Chiriqui, El Volcan, Panama. I have recently found
two specimens in the Banks collection in the Museum of Com-
parative Zoology from La Verbena, Costa Rica, (Tristan) which
appear to belong to this species.

Micrathena sexspinosa (Hahn), 1822

(Figures 174-180)

Epeira sexspinosa Hahn, 1822

E. furcata Hahn, 1822

Acrosoma sexspinosa Hahn, 1834

Plectana squamosa Walckenaer, 1841

A. obtusospinosa Keyserling, 1863

Keyserlingia coimigera O. P. -Cambridge, 1890. New synonymy.

A. calcaratum O. P.-Cambridge, 1890

A. obtusospinum Keyserling, 1892

A. calcaratum Keyserling, 1892

A. sedes Getaz, 1893

M . sexspinosa Simon, 1895

M. obtusospina F. P.-Cambridge, 1904

M. cornigera F. P.-Cambridge, 1904. New synonymy.

M. sedes F. P.-Cambridge, 1904

A. obtusispina Banks, 1909

A. sexspinosa Banks, 1909

M. obtusospinosa Petrunkevitch, 1911

M. sexspinosa Petrunkevitch, 1911

M . sedes Petrunkevitch, 1911

CHICKERING : MICRATIIENA IN CENTRAL AMERICA 457

M. cornigera Petrunkeviteh, 1911. New synonymy.

M. cornigera Beimoser, 1917. New synonymy.

M. sexspinosa Beimoser, 1917

M. cornigera Petrunkeviteh, 1925. New synonymy.

M. obtusispina Banks, 1929

M. cornigera Banks, 1929. New synonymy.

M. sexspinosa Petrunkeviteh, 1930

M. cornigera Boewer, 1942

M. sexspinosa Boewer, 1942

M. sexspinosa Kraus, 1955

M. cornigera Bonnet, 1957. New synonymy.

It now seems necessary to accept the complicated synonymy
the essentials of which are given above but I cannot regard M.
forcipata (Thorell) as a synonym for M. sexspinosa (Halm) as
Reimoser (1917), Roewer (1942), and Bonnet (1957) have done.
The Cuban specimens of M. forcipata (Thorell) are clearly dis-
tinct from M. sexspinosa (Ilahn), as I shall try to show in a
forthcoming paper. The specimens labelled M. sedes (Getaz) in
the Museum of Comparative Zoology are, in my opinion, all
immature females of M. sexspinosa (Halm). Immature females
have four pairs of abdominal spines and then, with the last
moult the pair between the surviving two pairs of dorsolateral
spines is often suppressed. Usually remains of these suppressed
spines can be found even in fully matured specimens. Banks
(1929) concluded that his A. cornigera 0. P. -Cambridge was
the male of M. sexspinosa but he did not formally synonymize it
with the latter. Other bibliographers have continued to keep
the two separate. Now I think we can be certain of the synonymy
as I have given it above.

Female hypotype. Total length 16.25 mm. The cephalic part
of the carapace is strongly raised ; the median fovea is a shallow
groove rather than a pit. The sternum is quite convex ; it bears
paired tubercles opposite coxae one to three and a posterior
tubercle extends between fourth coxae. The femora, especially
the first two, are well supplied with setigerous tubercles. The
features of the epigynum are shown in Figures 175-177. Abdo-
men : with three pairs of well developed spines and with another
pair represented by tubercles as shown in Figure 174. Color :
The carapace is a rich dark reddish brown with a whitish margin ;
dorsal ly the abdomen is a mosaic of whitish spots and streaks
intermixed with reddish and black spots and streaks; the inter-
mixture is quite variable but rather characteristic of the species.

458

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Male hypotype. Total length 5.59 mm. General form as shown
in Figure 178. Color: the dorsal surface is a mahogany brown
with light spots and light margin as indicated in the figure.
The most distinctive feature appears to be the long curved
process belonging to the tarsal hook ; this and other palpal fea-
tures are shown in Figures 179-180. The coxal hook and the
corresponding femoral ridge and groove are all lacking.

External Anatomy of Micrathena
Figures 174-180, M. sexspinosa

178

Fig. 174. Body of female; dorsal view.

Figs. 175-177. Epigynum from below, in posterior view, and in profile
from right side, respectively.

Fig. 178. Body of male ; dorsal view.
Fig. 179. Palpal tibia and tarsus of male.
Fig. 180. Basal tarsal hook.

CHICKEBING : MICRATHENA IN CENTRAL AMERICA

459

Collection records. The male and female hypotypes are from
Barro Colorado Island, C. Z., July and August, 1954, respec-
tively. The species is well known from many localities in Central
America, South America, and the West Indies. It appears to be
one of the two most abundant species on Barro Colorado Island.

Micrathena spinulata F. P. -Cambridge, 1904
(Figures 181-184)

If. spinulata Petrunkevitch, 1911
M. spinulata Reimoser, 1917
M. spinulata Roewer, 1942
M. spinulata Bonnet, 1957

This species seems to be extremely rare in collections. Ap-
parently it has not been recorded since the original specimens
were collected in Mexico and studied by the author of the
species. He had only females. During my period of work in the
British Museum (Natural History) in the summer of 1958 I
found the specimens studied by the author and identified the
individual used for F. P. -Cambridge's drawings. This speci-
men should be regarded as the lectotype and indicated as such.

181 i 184

External Anatomy of Micrathena

Figures 181-184, M. spinulata

Fig. 181. Abdomen of female; dorsal view.

Figs. 182-184. Epigynum from below, in posterior view, and in profile
from right side, respectively.

Lectotype. Total length 6.175 mm. Color pattern essentially
as given by the author of the species but I have noted consider-
able variation in the pattern among the various available speci-
mens. The median thoracic fovea is well defined together with

460

BILLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

three pairs of dorsolateral foveae. Sternum only slightly con-
vex; only slightly raised opposite the coxae. There are six ab-
dominal spines but all are hardly more than spinules (Pig. 181).
The epigynum is similar to that of M. gracilis (Pigs. 182-18-1).
The species is still only known from Mexico and, as far as I have
been able to learn, only from the original collections.

Micrathena striata F. P. -Cambridge, 1904
(Figures 185-188)

M. striata Petrunkeviteh, 1911
M. striata Reimoser, 1917
M. striata Eoewer, 1942
.1/. striata Bonnet, 1957

This is another very rare species in collections. My only op-
portunity to study it also came during my period of work in the
British Museum (Natural History) in the summer of 1958. It
seems highly probable that I was able to identify the specimen

185

External Anatomy of Micrathena

Figures 185-188, M. striata

Fig. 185. Abdomen of female; dorsal view.

Figs. 186-188. Epigynum from below, in posterior view, and in profile
from right side, respectively.

CHICKERING : MICRATHENA IN CENTRAL AMERICA 461

which the author of the species used as a basis for his description
and I have indicated that this should be designated the lectotype.
Numerous specimens from Guatemala are available for study.
The total length is about 8 mm. The carapace has a well defined
central fovea but no dorsolateral foveae. The sternum is essen-
tially like that of M. spinulata. The abdominal spines and gen-
eral form of the abdomen are shown in Figure 185. The essen-
tial features of the epigynum are shown in Figures 186-188. The
color of the abdomen is yellow with a series of narrow black
lateral lines. The species is apparently known only from Guate-
mala and only from the female.

MlCRATHENA SUBFLAVA Sp. nOV.

(Figures 189-193)

Female kolotype. Total length from AME to posterior border
of abdomen 8.515 mm. Carapace considerably overlapped by
abdomen; about 2.925 mm. long; 2.405 mm. wide opposite inter-
val between second and third coxae where it is widest ; strongly
gibbous just behind the well defined median fovea ; without well
defined dorsolateral foveae.

Eyes. Eight in two rows as usual ; viewed from above, both
rows moderately recurved ; viewed from in front, anterior row
gently recurved, posterior row gently procurved, all measured
by centers. Central ocular quadrangle slightly wider behind
than in front, slightly wider behind than long. Ratio of eyes
AME : ALE : PME : PLE = 10 : 8.5 : 12 : 8. AME separated
from one another by four-fifths of their diameter, from ALE by
four times their diameter. PME separated from one another by
five-sixths of their diameter, from PLE by ten-thirds of their
diameter. Height of clypeus equal to three-fifths of the diameter
of AME.

Chelicerae. Robust, quite gibbous in front; typical of females
in the genus. With a well defined fang groove having four teeth
along the promargin and three along the retromargin (some
variation noted between right and left sides).

Maxillae and Lip. Quite typical of the genus ; details regarded
as uimoteworthy for adequate description of the species.

Sternum. Moderately convex; with seven tubercles as com-
monly occur in the genus ; continued between fourth coxae which
are separated by about three-fourths of their width.

462

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Legs. 4123. Width of first patella at "knee" .29241 mm.,
tibial index of first leg 10. Width of fourth patella at "knee"
.30324 mm., tibial index of fourth leg 12.

Femora Patellae Tibiae Metatarsi
(All measurements in millimeters)

Tarsi

Totals

2.925

.975

1.950

1.625

.740

8.215

2.470

.942

1.570

1.495

.740

7.217

1.950

.650

.975

.910

.585

5.070

3.250

.845

1.755

1.950

.750

8.550

With many setigerous tubercles on legs but only a few spines
with details of spination regarded as unnoteworthy.

193

External Anatomy of Alicrathena

Figures 189-193, M. subflava

Fig. 189. Body of female; dorsal view.

Fig. 190. Spines at posterolateral angle of abdomen; lateral view.
Figs. 191-193. Epigynum from below, in posterior view, and in profile
from right side, respectively.

CHICKEBING : MICRATHENA IN CENTRAL AMERICA 463

Abdomen. General features as shown in Figures 189-190. Con-
siderably flattened dorsoventrally ; only slightly concave in mid-
dle of posterior dorsal region ; with nine pairs of short spines as
figured but the first pair of lateral marginal spines is often
reduced to blunt tubercles in paratypes. A prominent tubercle
or short robust spine lies on each side of the chitinous cone sur-
rounding the spinnerets.

Epigynum. The general characteristics of this organ are
shown in Figures 191-193.

Color in alcohol. Legs, in general, a dull brown with some
variations; sternum a dusky yellow; carapace yellowish with
fine brownish dots. Abdomen: main part of dorsum yellowish
with variations ; irregularly black along the lateral margins and
posterior border; lateral sides with narrow broken black stripes
alternating with yellowish stripes; region of epigynum and cone
around spinnerets reddish brown.

Type locality. Ilolotype female from Barro Colorado Island,
C. Z., August, 1950. Several paratype females from the same
locality: June, 1934; June-August, 1936; June, 1939; July-
August, 1954. The male is unknown.

Micrathena subspinosa F. P. -Cambridge, 1904
(Figures 194-197)

M. subspi7iosa Petrunkevitch, 1911
M. subspinosa Reimoser, 1917
M. subspinosa Roewer, 1942
M. subspinosa Bonnet, 1957

The correct treatment of this species is not at all clear. 1
have wavered between the decision to regard it as a variant of
M. duodccimspinosa (0. P. -Cambridge) and retaining it as a
valid species and have finally decided on the latter course, at
least for the present, i had an opportunity to study the type
in the British Museum (Natural History). The author of the
species gave the important features very briefly as follows :
"almost precisely similar to M. 12-spinosa in general character,
but the third, lower cusp on the posterior bifid spur is remote from
the two main cusps in the middle of the spur behind, and thus
becomes trifid. The posterior margin of the vulva, though pre-
senting a minute median angle, has not a tongue-like prominence
which is noticeable in 71/. 12-spinosa." Color: "similar to that
of M. 12-spinosa, the posterior spurs dark brown, apically black."

464

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

The length is about 6 mm. Figure 194 is intended to show rela-
tionships among the spines at the posterolateral angle of the
abdomen. Figures 195-197 are intended to show the chief fea-
tures of the epigynum. Known only from Guatemala and only
from the female.

External Anatomy of Micrathena
Figures 194-197, M. subspinosa

Fig. 194. Spines at posterolateral angle of abdomen ; lateral view.
Figs. 195-197. Epigynum from below, in posterior view, and in profile
from right side, respectively.

Micrathena trapa (Getaz), 1891

Acrosoma trapa Getaz, 1891
M. trapa F. P.-Cambridge, 1904
M. trapa Petrunkevitch, 1911
AI. trapa Eeimoser, 1917
M. trapa Eoewer, 1942
M. trapa Bonnet, 1957

F. P.-Cambridge (1904) did not include any description of
this species and did not include it in his keys. The brief de-
scription given by Reimoser (1917) was copied from the state-
ment given by the author of the species and gives little basis for
its recognition. It seems probable that the species exists among
others which are better known and well described. The male is
unknown. I have been obliged to omit the species from further
consideration in this paper.

Micrathena triserrata F. P.-Cambridge, 1904

(Figures 198-201)

Acrosoma triserrata Banks, 1909
M . triserrata Petrunkevitch, 1911
M. triserrata Eeimoser, 1917

CHICKERING : MICRATIIENA IN CENTRAL AMERICA

465

M. triserrata Koewer, 1942
.1/. triserrata Bonnet, 1957

Specimens in the Pickard-Cambridge collection from Guate-
mala and Costa Rica have been studied and a lectotype selected.
The following facts are taken from the lectotype in the British
Museum (Natural History) : Total length from AME to pos-
terior end of the triserrated abdominal fork 8.13 mm. Carapace
with a well defined central fovea behind which is a marked gib-
bosity ; there are also three pairs of dorsolateral foveae ; the head
is considerably raised and separated from the thoracic part by
diagonal grooves. The sternum is only slightly convex and
somewhat granulose. In addition to the abdominal spines shown
in F. P. -Cambridge's figures and named in bis description is,
apparently, a small lateral marginal spine at base of the fork
on each side (Pig. 198), but it appears to be somewhat variable.

External Anatomy of Micrathena

Figures 198-201, M. triserrata

Fig. 198. Abdomen of female; dorsal view.

Figs. 199-201. Epigynum from below, in posterior view, and in profile
from right side, respectively.

On the lectotype this small spine is unilateral (possibly lost in
handling), barely represented in one paratype, and lacking on
the third specimen. Features of the epigynum are shown in
Figures 199-201. The male is unknown. The species is known
only from Guatemala and Costa Rica.

460 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Micrathena uncata F. P. -Cambridge, 1904

(Figures 202-205)

.)/. uncata Petruiikevitch, 1911
M. uncata Reimoser, 1917
M. uncata Roewer, 1942
M. uncata Bonnet, 1957

This is another species which appears in collections only very
rarely. Apparently it has not been reported until the present
time since its first appearance in the collections studied by F. P.-
Cambridge (1904). I appear to have two specimens which belong
here. Only males are known.

Male hypotype. Total length 4.5 mm. The general appearance
(Fig. 202) is similar to that of 31. parallela (O. P. -Cambridge).
The first two pairs of tibiae do not have the modified short
ventral spines recorded by F. P. -Cambridge ; the first pair of
femora has a series of ventral and prolateral spines probably
modified for clasping (Fig. 203) ; the second pair of femora
seems to be devoid of these ; the ventral hook is lacking on the
first coxa and the corresponding ridge and groove on the second
femur are also lacking. The main features of the palpal tarsus
are shown in Figures 201-205. Color: Carapace with a broad,
median, light brownish stripe and a broad brown stripe on each
side (Fig. 202) ; the abdomen is light yellowish dorsally and
with numerous irregular white flecks and small indications of
black spots at anterior and posterolateral corners.

Collection records. F. P. -Cambridge had this species only from
Guatemala. The hypotype and one other specimen are from
Barro Colorado Island, C. Z., August, 1939.

Micrathena vitiosa (O. P. -Cambridge, 1890)

(Figures 206-210)

Acrosoma vitiosum O. P. -Cambridge, 1890

A. vitiosum Keyserling, 1892

.1/. vitiosa F. P.-Cambridge, 1904

.1/. vitiosa Petruiikevitch, 1911

M. vitiosa Reimoser, 1917

M. vitiosa Roewer, 1942

.1/. vitiosa Bonnet, 1957

While working in the British Museum (Natural History) in
the summer of 1958 I had my only opportunity to study this

CHICKERING : MICRATIIENA IN CENTRAL AMERICA

467

203

210

External Anatomy of Miorathena

Figures 202-205, M. uncata

Figures 206-210, M. vitiosa

Fig. 202. Dorsal view of body of male.
Fig. 203. Left first femur; ventral and prolateral spines.
Fig. 204. Left palpal tarsus.
Fig. 205. Another view of basal tarsal hook.
Fig. 206. Dorsal view of body of male (from O. P.-C).
Figs. 207-208. First and second femora, respectively; ventral views.
Figs. 209-210. Eight palpal patella, tibia, and tarsus; 209 especially to
show form of basal tarsal hook.

species. I found two specimens in the Cambridge collection and
one of these should be selected as the lectotype. The Pickard-
Cambridges had the species from Panama and Keyserling

468

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

(1892) reported it from Guatemala but I did not find these
specimens in the Keyserling collection. The total length is about
4.5 mm. The general form of the body is shown in Figure 206.
The first and second tibiae appeared to be spined in what
may be termed the usual manner. The first and second femora,
however, seemed to have special ventral spines as shown in
Figures 207-208. The third femur also has a row of spur-like
ventral spines. The most important features of the palp are
shown in Figures 209-210. The tarsal hook is particularly distinc-
tive. There is no ventral hook on the first coxa nor any ridge or
groove on the second femur. The female is completely unknown.

Micrathena zilchi Kraus, 1955

(Figures 211-213)

The holotype and several mature paratypes as well as several
more immature specimens were reported by Dr. Kraus in 1955
from El Salvador. I know of no mature specimens of this species
in American collections. I have one immature female from Hon-
duras which I am tentatively assigning to this species. Only
females are known. Some of the distinctive features of the
species, taken directly from the original description, may be

21 1

213

2 12

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: '■*■''■'.

*;■„" •

:V- •

-■ •*■ TV

W-i

®?

External Anatomy of Micrathena

Figures 211-213, M. zilchi

Fig. 211. Dorsal new of abdomen of female.

Figs. 212-213. Epigynum from below and in profile, respectively.

CIIICKERING : MICRATHENA IN CENTRAL AMERICA 469

given as follows : Total length 14 mm. ; central ocular quadrangle
almost square ; PME larger than AME ; general shape of abdo-
men and spination as shown in Figure 211; the epigynum ap-
pears as shown in Figures 212-213 ; the color of the abdomen is
bright, reddish yellow. The author of the species regards it as
closely related to M. xanthcpyga Simon, 1895 from Venezuela.
The accompanying figures have been copied directly from those
furnished with the original description.

SELECTED BIBLIOGRAPHY

Banks, Nathan

1898. Arachnida from Baja California and other parts of Mexico.
Proc. California Acad., Sci., Ser. 3, Zoology, 1(7): 205-309,
5 pis.
1909. Arachnida from Costa Rica. Proc. Acad. Nat. Sci. Philadelphia,

April, 1909: 194-234.
1929. Spiders from Panama. Bull. Mus. Comp. Zool., 69: 53-96, 4 pis.
Bonnet, Pierre

1957. Bibliographia Araneorum. 2. 3me partie. Toulouse: Les Artisans
de l'Imprimerie Douladoure.
Bryant, Elizabeth B.

1940. Cuban spiders in the Museum of Comparative Zoology. Bull.
Mus. Comp. Zool., 86(7): 249-532, 22 pis.
Cambridge, O. P.- and F. P.-Cambridge

1889- Arachnida- Araneida. In: Biologia Centrali- Americana. Dulau
1905 & Co., London.
Chamberlin, R. V. and Wilton Ivie

1936. New spiders from Mexico and Panama. Bull. Univ. Utah, 27:
No. 5, Biol, series, 3. No. 5:3-103, 17 pis.
Chickering, Arthur M.

1931. Notes and studies on Arachnida. I. Araneae from the Lancetilla
Valley, Honduras, I. Pap. Michigan Acad. Sci. Arts Letters,
13: 249-252.
1960. Six new species of Micrathena (Araneae, Argiopidae) from
South America with notes on known species. Proc. Zool. Soc.
London, 135(1) : 1-25, 78 figs.
Keyserling, Graf E. von

1863. Beschreibung neuer und wenig bekannter Arten aus der Familie
Orbitelae. Verh. Zool. Bot, Ges. Wien, 15: 799-856, pis. 18-21.
1892. Die Spinnen Amerikas. 4. Epeiridae. Niirnberg. Baur & Raspe.
Kraus, Otto

1955. Spinnen aus El Salvador (Arachnoidea, Araneae). Abhand.
Senckenberg. Naturf orsch. Ges., 493 : 1-112, 12 pis.

4-70 bulletin: museum of comparative zoology

Petrunkevitch, Alexander

1911. A synonymic index-catalogue of spiders of North, Central, and

South America, etc. Bull. Amer. Mus. Nat. Hist., 29: 1-809.
192.i. Arachnida from Panama. Trans. Conn. Acad. Arts Sci., 27:

51-248.
1930. The spiders of Porto Rico. Pt. 2. Trans. Conn. Acad. Arts Sci.,
30: 159-355, 240 figs.
Reimoser, Eduard

1917. Die Spinnengattung Micrathena Sundevall. Verh. zool.-bot. Ges.
Wien, 67(%): 73-160.
Roewer, C. F.

1942. Katalog der Araneae. Vol. 1. Bremen.
Simon, Eugene

1892- Histoire Naturelle des Araignees. Deuxieme Edition. 2 Vols.
1903. Librarie Encyclopedique de Roret, Paris.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 125, No. 14

THE PROSCALOPINAE, A NEW SUBFAMILY OF
TALPID 1NSECTIV0RES

By Katherine M. Reed

With Two Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

November, 1961

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Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 125, No. 14

THE PROSCALOPINAE, A NEW SUBFAMILY OF
TALPID 1NSECT1V0RES

By Katherine M. Reed

With Two Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

November, 1961

Xo. 14 --The Proscalopinae, a new Subfamily of Talpid

Insectivores

By Catherine M. Reed

INTRODUCTION

In tlif course of work on the insectivores of the Split Rock
local fauna of Wyoming (Reed, 1960), I had an opportunity
to examine the known material of Proscalops and its relatives.
Matthew (1928, pp. 70-71) suggested in passing that the forms
known to him might constitute a new family, although he con-
sidered this conjectural. With the discovery of at least three
more of these highly distinctive forms since Matthew's time, the
evidence now available indicates that at least a new subfamily of
the Talpidae must be erected for the group.

I am indebted to the authorities of the American Museum
of Natural History, the Chicago Natural History Museum, the
Kansas University Museum of Paleontology, and Amherst Col-
lege for the loan of specimens. Dr. Charles A. Reed, Dr. William
Turnbull, Dr. Reid Macdonald, Dr. Paul O. McGrew and Dr.
Raymond Alf all kindly assisted me with information about
material in their collections. I also wish to thank Prof. Bryan
Patterson and Mi-. Craig C. Black for their help in this study,
and the Mammal Department of the Museum of Comparative
Zoology for access to Recent talpid and chrysochlorid material.
1 am also grateful to Prof. Harry B. Whittington for his assist-
ance with the photography.

The following abbreviations are used :

A.C.M., Amherst College Museum; A.M.N.H., American Mu-
seum of Natural History; C.N.H.M., Chicago Natural History
Museum; K.U.M.V.P., Kansas University Museum of Vertebrate
Paleontology; M.C.Z., Museum of Comparative Zoology; 1.,
length, antero-posterior ; w., width, transverse; trig., trigonid;
tab, talonid.

TAXONOMY

Family TALPIDAE

PROSCALOPINAE, subfam. nov.

Type genus, f'roscah/ps Matthew, 1901.

Diagnosis. Skull chrysochlorid-like ; dentition of talpid type.

474 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Skull wide and deep in posterior part ; flexure in maxillary,
between P3 and P4, stronger than in Nctirotrichus; lateral pro-
jections on premaxillaries; slight sagittal and occipital crests;
palate long relative to skull length ; antorbital rim confined to
maxillary, not joining main body of zygoma. Wide lingual shelf
on upper molars with hypocone varyinglv developed; dental
formula [3 C* P3 M?. .

11! o

Included genera. Proscalops Matthew, !!)()! ; Mesoscalops
Reed, 1960; Oligoscalops gen. nov.

Range. Mid-Oligocene to mid-Miocene, North America.

Only Proscalops miocaenus and Mesoscalops scopelotemos have
hitherto been clearly diagnosed. Diagnoses of all species, in
order of their chronologic appearance follow.

Oligoscalops gen. nov.

Type species. Oligoscalops whitmanensis sp. nov.

Range. Mid-Oligocene, Wyoming and Colorado.

Diagnosis. Roots of P3 partially divided, tooth laterally
compressed, abutting against P1 ; P4 with large parastylar area
and lingual cusp situated at mid-line ; protocones of molars di-
rected antero-internally ; M2 wider than long; metastyle of M3
subparallel to line joining protocone and metacone ; hypocone
rudimentary.

Oligoscalops whitmanensis sp. nov.

Type. C.N.H.M. no. P 25800, partial skull and jaws.

Hypodigm. Type and K.U.M.V.P. no. 814-'], left ramus with
P2-M3.

Horizon and locality. Mid-Oligocene, type from Brule fm.,
Whitman, Niobrara County, Wyoming, collected by Dr. Paul
0. McGrew. K.U.M.V.P. no. 8143 from middle Cedar Creek mem-
ber of White River fm., W 1/2 sec. 7, T 11 N, R 53 W, Logan
County, Colorado, collected by Dr. Edwin C. Galbreath.

Diagnosis. As for the genus; the smallest known member of
the Prosealopinae.

reed: proscalopinae 475

Measurements (in millimeters).

C.N.H.M. no. P 25800

M1 left 1.2.2

M3 left 1. 1.4

w. 1.7

w. 1.4

right 1. 2.1-

right 1. 1.4

w. 1.65

w. —

M- left 1. 1.7

M2 right 1. 1.95

w. 2.0

w. trig. 1.35

right 1. 1.8

w. tal. 1.35

w. 2.1

Skull width at M2 : 9.0

K.U.M.V.P. no. 8143

Ml left 1 .-

M3 left 1. 1.5

w. tal. 1.0

w. trig. 1.3

M2 left 1. 2.1

w. tal. 0.9

w. trig. 1.7

w. tal. 2.1

PEOSCALOPS Matthew

Type species. Proscalops miocaenus Matthew.

Range. Late Mid-Oligocene to early Miocene.

Emended diagnosis. Lateral premaxillary projections dorso-
ventrally compressed; diastema between P3 and P4 ; P3 small,
oval, single rooted; P4 lacking parastylar area with lingual cusp
anterior to mid-line ; upper molars with or without well developed
hypocones; parastyles incipient on M1, definite on M2.

Proscalops miocaenus Matthew

Matthew, 1901, pp. 375-376, figs. 1-2; 1909, pi. 49, fig. 5; Gal-
breath, 1953, p. 49.
2 MCZ (Reed 6007 Mre7 Aug7

Type. A.M.N.H. no. 8949a, broken skull and jaw.

Hypodigm. Type only.

Horizon and locality. Mid-Oligocene, Vista beds, Cedar Creek,
Logan County, Colorado, collected by Dr. W. D. Matthew.

Emended diagnosis. Smallest known species of the genus. P4
relatively narrower than in Proscalops tcrtius, lingual cusp di-
rected posteriorly ; lingual portions of upper molars narrower
than in other species of the genus, with hypocones and proto-
styles less developed. M3 compressed antero-posteriorly, lingual
portion narrow, hypoeone rudimentary.

476 BULLETIN' : MUSEUM OF COMPARATIVE ZOOLOGY

Measurements (in millimeters).
A.M.N.H. no. 8949a

P4 left 1.2.0 M2 right I. 2.2

w. 1.7 w. 2.7

right 1. 2.0 M3 right 1. 1.5

w. 1.8 w. 1.8

M1 right 1. 2.8

w. 2.5

Palatal width at M2 : 9.4 (from Matthew, 1901, p. 376").

Proscalops tertius sp. now
Type. A.M.N.H. no. 19420. partial skull and left ramus with

Mx-3-

Hypodigm. Type only.

Horizon and locality. Oligocene, "White River fin.," possibly
Brule, "Badlands, South Dakota," collected by Dr. G. L. Jep-
sen. No more precise information is available.

Diagnosis. Teeth very slightly larger and hypocones better
developed than in Proscalops secundus; lingual cusp of P4 essen-
tially conical, larger than in /'. miocaenus, no part extending
anterior to labial cusp; metaconid of M2 with minute metastylid.
wide opening of talonid valley to interior as in Mogera wogura.

Measurements (in millimeters).
A.M.N.H. no. 19420

P4 left 1. 2.1

M3 left 1. 2.4

w. 2.1

w. -

right 1. 2.1

w. 2.2

w. 2.1

M1 left 1. 3.2

Al, left 1.2.85

w. 2.7

w. trig. 1.6

right 1. 3.2

w. tal. 2.05

w. 2.8

M2 left 1. 2.7

M2 left 1.2.8

w. trig. 1.8

w. 2.65

w. tal. 2.1

right 1. 2.9

M3 left 1. 2.2

w. 2.75

w. trig. 1.6

w. tal. 1.3

Skull width at M2 : 12.0

REED: PROSCALOPINAE 477

Proscalops secundus Matthew

Matthew, 1909, p. 559, pi. 51, tigs. 3, 4; Galbreath 1953, p. 49.

This species has never been formally described or diagnosed.
As Galbreath points out (1953, p. 49, footnote) "the type desig-
nation and specific name must be cited as figures 3 and 4 of plate
51, and the accompanying legends on page 559 of 'The Carnivora
and Insectivora of the Bridger Basin Middle Eocene' (Matthew,
W. D., 1909, Am. Mus. Nat. Hist. Mem., vol. 9, pt. 6)."

Type. A.M.N.H. no. 13798.

Hypodigm. Type only.

Horizon and locality. Early Miocene, "lower Rosebud" of
Matthew, Bear-in-the-lodge Creek, South Dakota, collected by
Dr. W. D. Matthew.1

Emended diagnosis. Larger than P. miocaenus ; lingual cusp
of P4 broader and extending farther anteriorly than in P. mio-
caenus, about as in Mesoscalops scopelotemos ; lingual portion
of M13 broader, and hypocones better developed on M1"2 than
in P. miocaenus ; posterior root of zygomatic arch joining flange
extending up side of cranium.

Measurements (in millimeters).
A.M.N.H. no. 13798

P4 left 1. 1.85 M1 left 1. 3.1

w. 1. 1.6 w. 2.6

right 1. 1.9 right 1. 2.7

w. 1.6 w. 2.5

M2 left 1. 2.55 M3 left 1. 2.3

w. 2.8 w. 1.9

right 1. 2.4 right 1. 2.2

w. 2.8 w. 1.8

Skull width at M2 : 10.7; skull is slightly crushed transversely.

Mesoscalops Reed

Type species. Mesoscalops scopelotemos Reed 1960.

Range. Mid-Miocene, Wyoming.

Diagnosis. In general similar to Proscalops, but differing as
follows : protocone of upper molars more rounded and somewhat
larger than hypocone, situated more nearly directly internal to

i Macdonald (pers. eomm.) suggests that the horizon may In- either Monroe
Civek formation or Harrison formation.

478 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

paraeone, imparting a square appearance to the outline of teeth ;
P4 with lingual face faintly grooved and with wide lingual
shelf; lower molars with small median eingulum just above gum
line between hypo- and protoconids, talonid of Mi wider than
trie'onid.

Mescoscalops scopelotemos Reed

Reed, 1960, pp. 2-4, 7-8, pi. 1, pi. 2, figs. 1, 2.

Type. A.C.M. no. 10461, isolated left M1.

Hypodigm. Type and numerous isolated teeth including
P4-M\ Mj-o. (See Reed 11)60. p. 2).

Horizon and locality. Mid-Miocene; NW 1/4 sec. 36, T 29 X,
R 90 W, Fremont County, Wyoming, from the vicinity of the
Brachycrus quarry seven miles west of Muddy Gap filling sta-
tion, in a draw about 1/4 mile south of U.S. highway 287.

Diagnosis. As for the genus.

Measurements. See Reed 1960, pp. 7-8.

MORPHOLOGY

The subfamily Proscalopinae is at present represented only
by skulls and partial rami or by isolated teeth. Leaving aside for
the moment the possible but questionable association of Arctory-
ctes-Cryptoryctcs with these forms, there are no postcranial ele-
ments known to me that can definitely be assigned to the species.

The skull

No sutures can be detected in any of the specimens, a situation
not uncommon among small fossorial forms. The anterior por-
tion of tbe skull is preserved only in Proscalops seciindus and
Oligoscalops whitmanensis. Tin- two are similar in having lat-
eral projections on the premaxillaries. In Oligoscalops these pro-
jections are abraded and now extend to a point above the roots
of I1 ; it is unlikely that they extended farther forward. The
lateral projections are only faintly demarcated from the dorsal
surface of the snout ; a slight groove partially separates the
nasals and the projections posteriorly. The lateral projections
in P. secundus are more distinct than in the earlier form. As
preserved, they extend slightly beyond I1. The processes form
conspicuous projecting shelves, fiat dorsally, situated well be-
low the level of the nasals. The edges of the projections tip

reed: proscalopinae 47!)

upward very slightly. These projections are distinctly different
from any structures in the same area in other talpids. There
are slight dorsal swellings in the premaxillaries of the various
talpids examined -- Scapanus, Parascalops, Talpa, Neurotrichus,
Condylura, Scalopus, Mogera. Uropsilus, Desmana, Galemys,
Bhynconax — but in no ease would these swellings significantly
change the round or squarish outline of a transverse section
through the anterior pari of the skull. A similar section through
either of the fossil skulls would give an elliptical section with
the major axis horizontal. The projections are more nearly com-
parable to the premaxillary projections of the chrysochlorids, but
here too there are distinct differences. The narial opening of the
Proscalopinae is situated immediately above the incisors with
no antero-dorsally inclined bony shelf above I1, such as occurs in
the African chrysochlorids, especially in Chlorotalpa. In Anibly-
somus and Eremitalpa the lateral projections are on the same
level as the nasals and tend to converge ventro-medially, rather
than more medially as in the Proscalopinae. In all chrysochlorids
the projections extend beyond the tooth-bearing portion of the
premaxillary, and in none are the projections as distinctly dif-
ferentiated from the general outline of the skull as they are in
P. secundus. Oligoscalops more nearly resembles the chryso-
chlorids in this latter respect. In a specimen of Eremitalpa
(M. C. Z. no. 39614), the cartilage of the rhinarium remains. The
cartilage is attached to the premaxillaries ventrally and the
nasals dorsally; it is supported ventro-laterally by the lateral
projections. It is likely that the projections in the Proscalopinae
served a similar purpose. The similarity in structure to the
chrysochlorids thus suggests a nasal region more chrysochlorid-
like than mole-like.

On the skull of Oligoscalops there are two slight depressions
above I3, one on either side. These are on the dorsal side of
the premaxillary and are slightly deeper antero-medially than
posteriorly. Analogous depressions are not immediately evident
on P. secundus, but may be represented by the dorsal surface of
the lateral projections and possibly the vertical portions of the
premaxillary medial to the projections. Nothing similar occurs
either in talpids or in chrysochlorids.

When the skulls of Proscalops secundus, Oligoscalops wliit-
manensis, and Proscalops tertius are viewed in profile, a feature
common to all can be seen, namely, a notable difference in the

480 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

depth anteriorly and posteriorly. This difference can be meas-
ured only in P. secundus and tertius, where the cranial region
is reasonably complete, but a similar difference is clearly sug-
gested in Oligoscalops. The difference between the two dimen-
sions is above or at the upper limit of similar measurements in
both talpids and chrysochlorids.

The tooth row in the Proscalopinae shows a flexure or arching
between P3 and P4, even more pronounced than in Neurotrichus.
Anterior to P4, the tooth row tends to parallel the dorsal profile
of the antorbital portion of the skull. Posterior to P4 the tooth
row and the skull profile diverge, the tooth row descending.
This characteristic can be seen in all the skulls, even in that of
the poorly preserved P. miocacnus; it is best shown by P.
secundus. This flexure occurs at the shallowest part of the skull.
A similar flexure cannot be found in other talpids except Neuro-
trichus, or in the chrysochlorids.

The structure of the zygomatic arch distinguishes the Proscalo-
pinae both from other talpids and from the chrysochlorids. The
anterior part of the zygoma is preserved in all the available
skulls. The anterior part of the arch in all is slender and
rounded, stouter than in Recent talpids, but not as deep as in the
chrysochlorids. The entire arch is preserved on the left side of
P. miocacnus ; it maintains its diameter throughout as in modern
talpids, the cross-section of the posterior root being essentially
the same as that of the anterior. Tn P. tertius, the left side of
the skull preserves a small rounded posterior root ; the right side
corresponds, although badly worn. P. secundus likewise preserves
a small rounded posterior root. This is a notable difference from
chrysochlorids in which the arch increases greatly in depth pos-
teriorly. The arch in the fossil forms shows no upward bending
as in Uropsilus or Rliijnconax. The facial regions of P. secundus,
P. tertius and Oligoscalops are widest at the level of M2. The
zygomatic arch leaves the side of the facial region above the pos-
terior margin of M2, as in Scapanus californicus and Uropsilus.
In all other talpids, the arch leaves opposite M3 or the posterior
margin of the tooth row. In chrysochlorids the arch begins above
M2, except in Ercmitalpa. The antorbital border, which, due to
the size of the infra-orbital foramen, forms a bar as in other
insectivores, does not reach to the zygoma but joins the main body
of the maxillary medial to the zygoma. The bar has a very
slight posterior slant to it. This feature of the skull is in decided
contrast to the Talpinae, Condylurinae and Sealopinae and to the
chrysochlorids in which not only does the bar join the zygoma.

reed: proscalopinae 481

but it also has a distinct posterior slant, uniting with the arch
posterior to the tooth row. In Mogera, the bar appears to be
attached to the side of the facial region, but here the posterior
slant is greater than in the Proscalopinae. In Uropsilus, however,
the bar is stout, posterior to the infraorbital and the large lac-
rymal foramina and joins the maxillary. It does not make up
any part of the zygoma. The bar in Desmana joins medial to the
rim of the zygoma, but is not attached to the maxillary.

In P. secundus, there is evidence of a squamosal flange extend
ing up the side of the skull, suggestive of the chrysochlorid con-
dition. However, the zygoma plays no part in the formation of
the flange in P. secundus as it does in the chrysochlorids. It is
impossible to determine the original width due to breakage.
Matthew's figure (1909) shows that some of this flange has been
lost since the photograph was made. The dorsal part of the flange
is lacking, but it is probable that it extended to the slight sagittal
crest present in this form. At a point approximately one-third
of the distance between the glenoid fossa and the sagittal crest
there is a buttress that divides the depression made by the flange
into two portions, the ventral evidently deeper than the dorsal.
A somewhat similar buttress is found in AmMysomus and
Eremitalpa at the point where the flange joins the side wall of
the cranium. However, the flange in P. secundus undoubtedly
did not end at the buttress but continued toward the sagittal
crest becoming progressively shallower dorsally. It is probable
that the squamosal in this form resembled that of the large chry-
sochlorid, Chrysospala.r, rather than the smaller forms, in the
degree of projection from the skull, as shown in Matthew's figure.
Desmana, which has both a sagittal crest and lateral, nearly hori-
zontal flanges at the extremities of the occipital crest, does not
in the least resemble /'. secundus in this region.

The occipital region of the Proscalopinae, best seen in P.
secundus, shows some resemblance to both talpids and chrysoch-
lorids. The fossil skull preserves a trace of the occipital crest
found in Recent talpids. especially prominent in Desmana. This
crest is not found in the chrysochlorids; instead there is a crest
which follows, essentially, the fronto-parietal suture above and
the junction of the squamosal and the posterior part of the
zygoma below. This crest is also present in the Proscalopinae.

The palate in the Proscalopinae extends posteriorly to a line
joining the posterior borders of M3, as in Neurotrichus. This
character easily distinguishes the Proscalopinae from Talpa,

482

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Mogera, Uropsilus, Desmana and Scalopus, in which the palate
ends well posterior to M3, or from Condylura in which the
palate ends anterior to M3. The ehrysochlorids resemhle Talpa,
etc., in this respect. The incisive foramen in P. secundus and
Oligoscalops is situated immediately medial to a point between
I1"2. The incisive foramen is more posterior in ehrysochlorids.
The posterior palatal foramina can he seen in P. secundus and
Oligoscalops; they occur on a line joining the posterodabial roots
of the first molars. There are no palatal vacuities. Although
neither the skull of P. secundus nor that of Oligoscalops is com-
plete or free from distortion, it is obvious on inspection that the
palate is longer relative to the total skull length and to basi-
cranial length than in Kecent talpids or in chrvsochlorids (Table
I)-

Table

Length of

palate

basicranium

Genus

in cm.

Basicranium/Palate

Proscalops secundus

1.54

1.23

.79

Oligoscalops v

hitmanensis

1.14

1.0

.88

Scapanus

1.53

2.1

1.3

Scalopus

1.43

1.69

1.17

Condylura

1.32

1.95

1.4

Mogera

1.59

2.44

1.4

Neurotrichus

1.0

1.23

1.2

Parascalops

1.38

1.79

1.3

Talpa

1.33

1.96

1.4

Uropsilus

1.0

1.14

Desmana

3.20

2.56

1.25

Chrysospalax

1.74

1.67

.96-

Amblysomus

1.22

1.23

1.00

Eremitalpa

0.87

1.0

1.1

Chlorotalpa

0.86

1.23

1.4

The condylar region of most Recent talpids easily distinguishes
them from the Proscalopinae. With the exception of Mogera,
the condyle is neither as stout nor as ventrally inclined a struc-
ture as in the fossil forms. Recent talpids also possess a small
depression at the anterior end of the condyle, a feature not seen
in the Proscalopinae. Chrvsochlorids resemble the Proscalopinae

reed: proscalopinae 483

in the structures of this region. The foramen magnum in Recent
talpids has a notable nick that extends anteriorly to the level of
the posterior lacerate foramen or even slightly anterior to it.
The corresponding- nick in the Proscalopinae extends to a similar
position but is much less pronounced, due to the more ventral
position of the condyle. The same is true of the chrysochlorids.
The condylar and posterior lacerate foramina in the Proscalo-
pinae are situated close to the condyle ; the former is a narrow
slit just above the articular area of the condyle. The position of
these foramina is similar to that in Recent talpids. The carotid
foramen is closer and more nearly lateral to the condylar and
posterior lacerate foramina in the fossils than in Scalopus and
other Recent talpids, but is not greatly different in position from
that of the chrysochlorids. The carotid artery seems to enter the
posterior wall of the bulla as in the talpids. It is the most con-
spicuous foramen in the skull.

The bullae of P. tertius are irregularly quadrangular in out-
line and are neither as inflated as in the chrysochlorids nor as
flattened as in Recent talpids. They bear rounded crests that
run medially from the external auditory meatus, the anterior
and posterior parts of the bullae sloping upwards from the
crests. In Oligoscalops the bullae are less differentiated from
the skull, and the crests are less salient. No sutures between the
various elements of the bullae can be seen, but a basisphenoid
component was certainly present. As in Recent talpids, the
bullae merge anteriorly with the side walls of the posterior narial
passage but, in contrast to most of them, the basisphenoid and
elements anterior to it are little if at all inflated or cancellous
in the Proscalopinae. The basicranium in this area bears a slight
but notable median crest that runs forward to about the level of
the Eustachian openings. A similar but smaller crest can be
seen in Scalopus, and in Dcsmana this crest is slightly larger.
The external auditory meatus in /'. tertius is a large opening,
larger than in Scalopus. It is broken in the other fossil specimens.
The foramen ovale can be seen in P. secundus in the usual posi-
tion. Its large size suggests that it also gave passage to the
tendon of M. tensor tympani, as in the chrysochlorids and smaller
moles. It may also have housed a vein. The limits of the foramen
rotundum cannot be determined, but it is anterior to the fora-
men ovale and in the usual position. The Eustachian openings
are rather large and oval shaped, directed almost immediately
ventrally. The openings are larger than any observed in chry-
sochlorids. Placed slightly lateral and posterior to the external

484 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

auditory meatus is the stylomastoid foramen, and postero-medial
to it is the liyoidean vagina. The two are separated from the
external auditory meatus by a low bar of bone. Conditions here
are much as in the chrysoehlorids. In Recent talpids, such as
Scalopus, the openings arc situated postero-lateral to the carotid
foramen, not antero-lateral to it as in the fossils. The hyoidean
vagina is much more pronounced than in the Recent forms.
There arc two other noticeable depressions on either side of the
skulls, postero-lateral to the carotid foramen. One is almost di-
rectly posterior to the hyoidean vagina, the other posterior to this
but more medial. The latter depression is somewhat elongate
transversely. The function of these depressions is unknown.

The glenoid fossa of P. secundus and P. tertius is a much more
pronounced structure than in Recent talpids. It is wider trans-
versely and deeper, more as in chrysoehlorids. It differs from
these forms in having a prominent, anteriorly placed dorsal lip.
The postglenoid foramen is extremely small and is situated high
on the posterior face of the glenoid fossa near its lateral extrem-
ity. It is somewhat less lateral in position in the chrysoehlorids,
where it is on the medial face of the fossa and more nearly dorsal
to the external auditory meatus. The usual condition in Recent
talpids is for the foramen to be situated ventral to the posterior
wall of the fossa, immediately dorsal to the anterior cms of the
tympanic.

Fig. I. Diagram of the basieranial region of Prosealops tertius. Stippled
areas represent broken parts of skull. x3. Abbreviations: e.f., carotid fora-
men; co.f., condylar foramen; Eu., Eustachian opening; h.v., hyoidean
vagina; p.l.f., posterior lacerate foramen; s.f., stylomastoid foramen.

reed: proscalopinae 485

The lower jaw

Rami are known in Proscalops tertius, P. miocaenus and Oligo-
scalops. These are slender and relatively uniform in depth. The
rami of Oligoscalops show a slight bowing down below P^M^,
corresponding to the flexure in the upper tooth row. This bowing
down is unlike anything seen in other talpids or in chryso-
chlorids. The symphysis was evidently elongate but weak. This
is suggested by the rugose area on one of the rami of Oligoscalops,
which extends posteriorly to P4. In this respect the Proscalo-
pinae do not differ from most talpids. The mental foramen, a
single opening, is preserved only in Oligoscalops, where it is
situated below P3.

The posterior portion of the ramus is most completely pre-
served in P. miocaenus and the description is based on this speci-
men. The coronoid process rises at right angles to the main body
of the ramus and, although incomplete at the tip, does not seem
to curve posteriorly. It is not as narrow as in Recent talpids,
and the posterior border, as in Mogera, has a forward slope.
The condyle is wider than in Recent forms and is situated on
a long stout neck. The angular process is short, stout and wide
and leaves the body of the ramus at a lower level than in Recent
talpids. In Oligoscalops, so far as can be told from the material,
it is possible that the angle may have left the ramus at a some-
what higher level. The angle is not dorso-ventrally compressed
as in Uropsilus. In comparison with the chrysochlorids, the pro-
scalopine ramus is more slender. In the African forms, with
the exception of the large Chrysospalax, the coronoid process is
greatly reduced. The condyle is similar in inclination and in
the structure of the neck, but the articular surface in the Prosca-
lopinae faces essentially posteriorly whereas this surface in the
chrysochlorids faces almost entirely dorsally. The angle of the
chrysochlorids differs in being deeper and more compressed as
well as projecting well below the horizontal ramus. The two
groups are very distinct in this region.

The dentition

The diagnostic characters of the dentition of the members of
the subfamily have been given in the taxonomic section. A more
detailed treatment of the dentition and a brief comparison with
Recent forms follow.

Upper dentition. In both Oligoscalops and Proscalops secundus
the first incisor is enlarged and is flanked by two other small
apparently conical incisors. I 3 is smaller than I 2 in P. secondus;
these teeth are broken and represented only by the roots in

■iSQ BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Oligoscalops. The canine is simple and conical in both speci-
mens. It is larger than I - :: in /'. stridulus and Larger than the
first premolar in Oligoscalops. All the incisors, canines and an-
terior premolars are separated from each other by diastemata.
The premolar series consists of three teeth,2 interpreted as P1, 3~4.
In Oligoscalops, 1>;! is laterally compressed, has a partially di-
vided root, and is in contact with P4. In /'. miocaenus the two
anterior premolars are badly worn. P3 is not laterally com-
pressed as in Oligoscalops and has one root. A diastema separates
P3 and P4 in all species of Proscalops. In P. secundus, P1,3 are
simple, conical and single-rooted teeth. P:: is present on the
left side of P. tertius and agrees in form with that of P. secundus.
These teeth are not known in Mesoscalops.

P4 differs in the several species. In Oligoscalops, although
worn, it consists of a large labial paracone with a parastylar area
and a small conical lingual cusp. The latter cusp has a slightly
pinched appearance and is directed posteriorly. In all the species
of Proscalops and in Mesoscalops, P4 lacks the parastylar area.
In P. miocaenus, both the tooth as a whole and the lingual cusp
are relatively larger than in Oligoscalops ; again, the lingual cusp
is directed posteriorly. P4 of /'. tertius is larger still and also has
the essentially conical lingual cusp directed posteriorly. A
change in P4 becomes evident in P. secundus. In this species the
lingual cusp has broadened and is more shelf -like in shape. It is
also directed more medially than posteriorly, although no part
of the lingual cusp extends anterior to the blade-like paracone. In
Mesoscalops the antero-posterior broadening of the lingual cusp
is carried further ; it is here distinctly shelf-like. The lingual
cusp extends slightly anterior to the paracolic, although still
directed essentially medially; it shows a partial division and the
margin bears irregularities in the form of swellings. These are
also present but are less numerous in P. secundus. One specimen
of Mesoscalops exhibits a small cuspule on the posterior side of
the labial cusp. In Oligoscalops and the species of Proscalops the
tooth has three roots, two labial and one lingual. In Mesoscalops
the roots have fused, but traces of the original divisions remain.

M1 in Oligoscalops is triangular in general outline, with the
protocone directed anteriorly. The hypocone is rudimentary, a
mere protuberance labial and posterior to the protocone. The
blade-like paracone is smaller than the Y-shaped metacone. In
P. miocaenus the tooth is generally similar, although the hypo-
cone is somewhat better developed and the lingual portion of

-Tin- premolar sn-ics might be I'1 --■ ' by analogy with Uropsilus.

reed: proscalopinae 487

the tooth therefore somewhat broader. A rudimentary protostyle
is present. Para- and nietaeones are similar to those of Oligosca-
lops. In /'. tertius the hypocone is well developed. There is a
small but distinct protostyle anterior to the protoeone, and a
minute hypostyle posterior to the hypocone. The protoeone is
directed somewhat more medially than in the earlier forms. The
paracone is again smaller ami more blade-like than the metacone.
/'. secundus continues the trend toward a squaring up of the
outline of the tooth. The lingual shelf is wider, and the hypocone
and protostyle better developed. The paracone is slightly more
Y-shaped than in earlier forms, and there is a small distinct
parastyle. A metastyle is also present. M1 of Mesoscalops has
the protoeone and hypocone more nearly equal in size, and lacks
a protostyle, which gives a distinctly square appearance to the
tooth. The para- and metacones resemble those of /'. tertius,
rather than those of P. secundus. Both the parastyles and meta-
styles are present.

M2 of Oligoscalops and /'. miocaenus are again similar, al-
though that of the latter is larger and has a more pronounced
protoeone and parastyle ; a metastyle is lacking. In both, the
paracone and metacone are V-shaped. P. tertius is similar in
general, again enlarging the lingual portion of the tooth by a
well developed hypocone. It also has a parastyle, but no distinct
metastyle. The tooth is essentially square, with paracones and
metacones subequal in size. P. secundus is very similar, possess-
ing a slightly more distinct parastyle and a rudimentary meta-
style. In Mesoscalops the protostyle is less distinct, due to the
equalization of the protoeone and hypocone. A parastyle is pres-
ent, but the metastyle is incorporated in the crest from the
metacone.

M3 in Oligoscalops and /'. miocaenus is distinctly triangular,
with the protoeone directly lingual to the paracone. The hypo-
cone is rudimentary in Oligoscalops, absent in /'. miocaenus.
There are no distinct stylar cusps in either form. This tooth, in
both, is antero-posteriorly compressed, slightly less so in Oligo-
scalops. The posterior margin of the tooth is formed by the blade-
like metacone. M3 of /'. tertius is similar although larger. The
hypocone, however, is clearly present and lingual to the metacone
which again forms the posterior margin of the tooth. A proto-
style is suggested by a slight swelling. /'. secundus has a wider
lingual area with a distinct hypocone and protostyle. The meta-
cone still forms the posterior margin of the tooth, although on
RM3 there is a minute cuspule posterior to the metacone. A

488 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

parastyle is present but there is no distinct metastyle. In Meso-
scalops the hypocone is small and rather variable in shape, and
the protostyle is very poorly developed. Xo distinct stylar cusps
are seen. In all the specimens, M3 is the smallest of the molars.
All the upper molars have three roots, two labial and one
lingual. In Mesoscalops the lingual root is stouter and longer
than the labial roots. The same is suggested for the others. In
all, the protocone acquires a decidedly pinched appearance with
wear, but the degree of pinching is not diagnostic.

Lower dentition. The incisors, canines and anterior premolars
in Oligoseolops, the only form in which these teeth are known, are
all broken or badly worn. The Kansas specimen suggests that
the canine and F1 and P3 were simple, conical and single rooted.
P4 is laterally compressed and two rooted. In M1 the talonid is
wider than the trigonid, judging from the fragments that remain,
and the opposite is true of M2. A small anterior cuspule is pre-
served at the base of the paraconid of RM2. In M3 the trigonid
is wider than the talonid. The relative sizes of the cusps on the
molars cannot be determined, due to breakage. Matthew (1909,
pi. 49) figures a right ramus of the type of P. miocaenus3 in
which the relative trigonid-talonid widths are as in Oligoscalops.
The metaconid of M2 is larger than either the paraconid or the
entoconid. An anterior lingual basal cuspule is present in M3.
P4 of this specimen has a small heel, but is essentially conical
and apparently two rooted. P3 is simple, conical and single
rooted. All the left lower molars of P. tertius are known. In Mx
the trigonid is narrower than the talonid. The paraconid and
metaconid diverge more than in M- ;\ and a small posterior
cingular cuspule and a slight swelling on the lingual face of the
paraconid are preserved. In M2 the trigonid is wider than the
talonid. The metaconid is stout with a rudimentary metastylid ;
there are both antero-internal and postero-internal cuspules. In
M3 the trigonid is again wider than the talonid. There is only
an antero-internal cuspule. The lower dentition of P. secundus
is unknown. Mx of Mesoscalops has the trigonid narrower than
the talonid, with the paraconid and metaconid widely divergent.
A small postero-internal cuspule is present. M2 has a metastylid,
even more distinct than that in 7*. tertius, as well as antero-
internal and postero-internal cuspules. The trigonid is equal in
width to the talonid. M3 has a simple metaconid and both antero-
and postero-internal cuspules. The trigonid is wider than the
talonid.

3 This part of the type was not included in the material sent to me for study.

reed: proscalopinae 489

Comparison with Recent forms

Although the teeth of the Proscalopinae are talpid-like, a brief
comparison with Recent forms is desirable. Most of the Recent
talpids have three incisors, the first enlarged, the second and
third decreasing in size. Condylura, however, has a large eanini-
form second incisor. Larger than either I1 or the canine. Urop-
silus, apparently with only two incisors, possesses an enlarged I2,
but it is lower than I1. In Scalopus the second and third incisors
are minute and placed very close together. The second incisor is
separated from the very large I1 by a large diastema in Desmana.
The canines in all but Talpa and Mogera, where they are large,
are only slightly larger than the second or third incisor or the first
premolar, as in the Proscalopinae. Scalopus and Condylura are
the only Recent forms with a reduced premolar series. In Crop-
silus P3 is either present or absent; where present it is greatly
reduced.4 P2 in this form is larger than both P1 and P3. P4
shows the greatest difference between the Recent and fossil forms.
Condylura, Parascalops, Mogera, Neurotrichus, and Talpa all
have a small basal cusp anterior to the paracolic The lingual
cusp in these forms is extremely small. The general structure of
P4 of Oligoscalops shows some similarity to these five genera.
Both the anterior basal cusp and the lingual cusp are essentially
lacking in Sea pan us and Scalopus. P4 of VropsUus is more
similar to most of the Proscalopinae than other talpids, but has
a slight anterior cingulum not unlike Desmana. The lingual
portion of the molars is narrower and lacks the hypocone develop-
ment of the later Proscalopinae in all except Neurotrichus, Para-
scalops, Uropsilus and the members of the Desmaninae. In these
forms the wider lingual portion does not show any pinching of
the protocone, and the paracolic and metaeone of M1 in Neuro-
trichus are much simpler than in the fossils. The position of the
protostyle in Desniana is very different than in the Proscalopinae.
In all Recent talpids, excepting Scalopus in which they are essen-
tially lacking, the stylar cusps are more distinct than in the fossils.
In the lower dentition, Talpa, Mogera, and Condylura have two-
rooted premolar teeth. Parascalops and Condylura are the only
Recent talpids that have metastylids on the molars. Although
antero- and postero-internal cuspnles are generally present, in
no case do they connect with cingula as in the Proscalopinae.

•* In three of four specimens examined, P3 was present, although it was most
greatly reduced in a male skull. The one skull lacking P3 was also a male
Cabrera's dental formula is thus incorrect and should read for the upper
dentition either I- ("i P* M3 or 13 Ci P3 M3, not 12 Ci P3 M3 as given (Cabrera,
1925, Genera Mammalium. vol. 2, Insectivora, Galeopithecia) .

490

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Cingula are usually lacking, except in Uropsilus and some of the
Desmaninae.

Further comparison between the Proscalopinae and Recent
talpids in the dentition seems unnecessary; the extinct forms
clearly differ from the living in various characters and combina-
tions of characters. No comparison with the zalambdodont chry-
sochlorids is required.

Arctoryctes-Cryptoryctes and the Proscalopinae

It has been suggested by Matthew (1928) and by Schlaikjer
(1933) that the humeri described as Arctoryctes may belong to
the Proscalops group. This suggestion must now be considered.

Two points tend to support such an association. First, both
skulls and humeri have talpid characters, yet neither are typ-
ically talpid. (The humeri, of course, are very different from
those of the chrysochlorids. ) Second, the published geologic and
geographic ranges of the Proscalopinae and the Arctoryctes group
overlap (see Table II).

Table II
Oligocene

Early

Middle

Late

Miocene
Early Middle

Cryptoryctes

kayi
Cryptoryctes

sp.?
Arctoryctes

terrenus
Arctoryctes

galbreathi
Arctoryctes

sp.?

Montana

Colorado*

Nebraska

Montana
Colorado

S. Dakota

Wyoming**

Oligoscalops

whitmanensis
Proscalops

niiocaenus
Proscalops

tertius
Proscalops

secundus
Mesoscalops

scopelotemos

Wyoming
Colorado

Colorado
S. Dakotat

S. Dakota

Wyoming

* R( corded, but lost in the field.
•• Species not identified. (Reed, C. A., pers. coinni., 1961).
t Exact horizon uncertain.

reed: proscalopinae

491

A comparison of the skull length/humeras length ratio has
been made for modern talpids, for a chrysoehlorid, and for
selected proscalopine skulls and arctoryctine humeri. In two
cases, skull material and humeri are known from the same horizon
and locality: Oligoscalops whitmanensis (K.U.M.V.P. no. 8143.)
and Arctoryctes galbreaihi (K.U.M.V.P. nos. 0837-9839); and
Mesoscalops scopelotemos and an unidentified species of Arctory-
ctes. In the first case, unfortunately, the two best preserved
humeri are broken, and the measurements, kindly supplied by
0. A. Reed, are only approximate. No measurements are avail-
able for the humeri found with Mesoscalops scopel&temos. There
is a fair correspondence in time in one other case, that of
Proscalops secundus and Arctoryctes terrenus; the former is
probably from a somewhat earlier horizon in the Miocene of
South Dakota than the latter. All other occurrences are rather
widely separated in time or space. There is no known proscalo-
pine that could correspond to the small Cryptoryctes kayi. The
ratios (Table III) of Oligoscalops whitmanensis/ Arctoryctes gal-
breaihi and Proscalops secundus/ Arctoryctes terrenus do not
differ greatly from those of certain talpids but do differ from
that of the only chrysoehlorid available for comparison. These
ratios at least suggest that association is not impossible.

specimen

Seapanus

Condylura

NTeurotrichus

Chrysochloria

stuhlmani

Oligoscalops

whitmanensis

skull

Arctoryctes

galbreathi

humerus

Proscalops

secundus

skull

Arctoryctes

terrenus

humerus

Table III

skull/humerus ratio

remarks

229%
245%
304%

180%

252-256%

245%

from Reed, 1954
< < a a

it it i (

(long skull)

based on esti-
mated humeri
lengths

slight
difference
in age

492 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Although not impossible, the association of the skulls and
humeri is, of course, very far from proven. A typical talpid
humerus is known from the mid-Oligocene Cedar Creek member
of Colorado (Galbreath 1953), the same deposit that has yielded
specimens of Oligoscalops whitmanensis and Arctoryctes gal-
breathi. The earliest non-proscalopine North American talpid
is Domninoidcs; two species of this genus are known from the
earlier Miocene, one of them from beds just below those contain-
ing Proscalops secundus (Macdonald, pers. comm.). I have seen
typical talpid humeri from a deposit that has yielded remains of
Domninoidcs ("Valentine formation," Fort Niobrara locality)
and strongly suspect that these humeri are referable to that
genus. It seems likely that the Cedar Creek talpid humerus rep-
resents some otherwise unrecorded member of the family, but
the possibility that it is referable to Oligoscalops should not be
ignored — nor can the possibility be ignored that the Arctoryctes-
Cryptoryctes humeri may be referable to some other group alto-
gether. Only the fortunate discovery of proscalopine cranial or
dental material in unmistakable association with arctoryctine
humeri will settle the question.

After this study was completed, Russell (I960) suggested that
the arctoryctine humeri may belong to Micropternodus and re-
lated forms, since a humerus, identified by C. A. Reed as Cryp-
toryctes, occurs at a locality that has yielded " Eentrogomphios"
(= Micropternodus). Furthermore, if the skull proportions of
Micropternodus are similar to those of talpids, the skull/humerus
ratio of Micropternodus and Cryptoryctes (248^ ) is much closer
to those of talpids than to those of epoicotherids or apternodon-
tines, again suggesting a possible relationship between Micropter-
nodus and Cryptoryctes.

This ratio is not very different from that obtained by compar-
ing proscalopine skulls and arctoryctine humeri. However, as the
evidence supporting the association of the humeri with either
the Proscalopinae or the Micropternodus group is no more than
circumstantial at best, it is not possible to draw any conclusions
at this time.

CONCLUSIONS

The morphological characters of the skull obviously unite the
proscalopine species and clearly separate them from other tal
pids. As regards intergroup relationships, it can be stated that
Oligoscalops, the earliest known form, can be easily distinguished

reed: proscalopinae

493

from the rest of the subfamily on the characters of P3 and P4,
although the overall cusp pattern and the structure of the molars
are not very different from Proscalops. Within the genus Pro-
scalops there arc certain trends in the evolution of the dentition
that may easily be seen. There is a tendency to broaden the
lingual portion of the upper molars and 1>4, with a general squar-
ing of the outline of these teeth. In the lower molars there is a
progressive complication of the teeth by the addition of antero-
and postero-internal cuspules and the development of a metasty-
lid. McsoscaJops, the latest known form, while similar in the
general pattern of the lower molars, has modified the upper
molars by eliminating the protostyles. P4 of this form differs
in having incipient division of the lingual cusp as well as fusion
of the roots. This genus could, however, have been derived from
a form not unlike P. secundus. The mutual relationships within
the subfamily appear to be as in Figure 2.

M e soscalops

Prosco/ops secundus

Prose o/ops m/ocoenus

Proscahps ferf/us

Ol/goscalops

Fig. -. Diagrammatic representation of the relationship of the known
members of the Prosealopinae.

The Proscalopinae are very different from all other talpids so
far as the characters of the skull are concerned. In this the group
differs more from the other talpid subfamilies than these sub-
families do from each other. This su^-ests that it might be more
proper to group all other moles, with the exception of the Des-
maninae, in one subfamily. Our ignorance of the family is such,
however, that I do not take this step. If the curious arctoryctine
humeri should prove to be referable to the proscalopines, the
combination of cranial and humeral characters would certainly
warrant the erection of a new family for the Proscalopinae, as
Matthew suspected.

494 BULLETIN' : MUSEUM OF COMPARATIVE ZOOLOGY

It has been shown that there is a remarkable parallelism
between the proscalopines and the ehrysoehlorids. This is par-
ticularly evident in the premaxillary region, the squamosal
flange, seen particularly in P. secundus, and the general structure
of the basicranial region. The fact that these peculiar fossorial
adaptations have arisen more than once in the Insectivora re-
moves some of the uniqueness of the ehrysoehlorids.

REFERENCES

Cabrera, A.

1925. Genera Mammalium. Vol. 2, Insectivora, Galeopithecia. Madrid,
232 p.

Galbreath, E. C.

1953. A contribution to the Tertiary geology and paleontology of
northeastern Colorado. Univ. Kansas, Paleontologieal Contribu-
tions, Vertebrata, Article 4, pp. 1-120.

Hough, J. and R. M. Alf

195G. Chadronian mammalian fauna from Nebraska. Jour. Paleontol-
ogy, vol. 30, pp. 132-140.
Matthew, W. D.

1901. Fossil mammals of the Tertiary of northeastern Colorado. Mem.

Anier. Mus. Nat. Hist., vol. 1, pt. 7, pp. 355-447.
1909. The Carnivora and Insectivora of the Bridger Basin Middle
Eocene. Mem. Anier. Mus. Nat. Hist., vol. 9, pt. 6, pp. 291-567.
1928. Xenotherium, an edentate. Jour. Mammalogy, vol. 9, pp. 70-71.
Reed, C. A.

1954. Some fossorial mammals from the Tertiary of western North
America. Jour. Paleontology, vol. 28, pp. 102-111.

1956. A new species of the fossorial mammal Arctoryetes from the
Oligocene of Colorado. Fieldiana, Geology, vol. 10, no. 24, pp.
305-311.
Reed, K. M.

1960. Insectivora of the Middle Miocene Split Rock local fauna,
Wyoming. Breviora, Mus. Comp. Zool., no. 116, pp. 1-11 .
Russell, I). A.

1960. A review of the Oligocene insectivore Micropternodus borealis.
Jour. Paleontology, vol. 34, no. 5, pp. 940-949.
Schlaikjer, E. M.

1933. Contributions to the stratigraphy and paleontology of the Goshen
Hole area, Wyoming. I. A detailed study of the structure and
relationships of a new zalambdodont insectivore from the Middle
Oligocene. Bull. Mus. Comp. Zool. Harvard Coll., vol. 76, pp.
1-27.

PLATES

Plate 1

Fig. 1. Oligoscalops ichitmanensis, C.N.H.M. P25800, ventral view
Fig. 2. 0. whitmanensis, dorsal view
Fig. 3. 0. whitmanensis, left profile

Figs 1-3 approx. 2.5x
Fig. 4. Pioscalops tertius, A.M.N.H. 19420, stereoscopic pair, crown view,

approx. 4.5x
Fig. 5. Oligoscalops ichitmanensis, K.U.M.V.P. 8143, stereoscopic pair,

crown view, approx. 4.5x
Fig. 6. Proscalops tertius, ventral view
Fig. 7. P. tertius, left profile

Figs. 6 and 7 approx. 2.5x

PLATE 1

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